Soil penetration by maize roots is negatively related to ethylene-induced thickening.

Radial expansion is a classic response of roots to a mechanical impedance that has generally been assumed to aid penetration. We analysed the response of maize nodal roots to impedance to test the hypothesis that radial expansion is not related to the ability of roots to cross a compacted soil layer. Genotypes varied in their ability to cross the compacted layer, and those with a steeper approach to the compacted layer or less radial expansion in the compacted layer were more likely to cross the layer and achieve greater depth. Root radial expansion was due to cortical cell size expansion, while cortical cell file number remained constant. Genotypes and nodal root classes that exhibited radial expansion in the compacted soil layer generally also thickened in response to exogenous ethylene in hydroponic culture, that is, radial expansion in response to ethylene was correlated with the thickening response to impedance in soil. We propose that ethylene insensitive roots, that is, those that do not thicken and can overcome impedance, have a competitive advantage under mechanically impeded conditions as they can maintain their elongation rates. We suggest that prolonged exposure to ethylene could function as a stop signal for axial root growth.

The helical motions of roots are linked to avoidance of particle forces in soil.

Limitation to root growth results from forces required to overcome soil resistance to deformation. The variations in individual particle forces affects root development and often deflects the growth trajectory. We have developed transparent soil and optical projection tomography microscopy systems where measurements of growth trajectory and particle forces can be acquired in a granular medium at a range of confining pressures. We developed image-processing pipelines to analyse patterns in root trajectories and a stochastic-mechanical theory to establish how root deflections relate to particle forces and thickening of the root. Root thickening compensates for the increase in mean particle forces but does not prevent deflections from 5% of most extreme individual particle forces causing root deflection. The magnitude of deflections increases with pressure but they assemble into helices of conserved wavelength in response linked to gravitropism. The study reveals mechanisms for the understanding of root growth in mechanically impeding soil conditions and provides insights relevant to breeding of drought-resistant crops.

Root anatomical traits contribute to deeper rooting of maize under compacted field conditions.

To better understand the role of root anatomy in regulating plant adaptation to soil mechanical impedance, 12 maize lines were evaluated in two soils with and without compaction treatments under field conditions. Penetrometer resistance was 1-2 MPa greater in the surface 30 cm of the compacted plots at a water content of 17-20% (v/v). Root thickening in response to compaction varied among genotypes and was negatively associated with rooting depth at one field site under non-compacted plots. Thickening was not associated with rooting depth on compacted plots. Genotypic variation in root anatomy was related to rooting depth. Deeper-rooting plants were associated with reduced cortical cell file number in combination with greater mid cortical cell area for node 3 roots. For node 4, roots with increased aerenchyma were deeper roots. A greater influence of anatomy on rooting depth was observed for the thinner root classes. We found no evidence that root thickening is related to deeper rooting in compacted soil; however, anatomical traits are important, especially for thinner root classes.

A Maize Inbred Exhibits Resistance Against Western Corn Rootwoorm, Diabrotica virgifera virgifera.

Insect resistance against root herbivores like the western corn rootworm (WCR, Diabrotica virgifera virgifera) is not well understood in non-transgenic maize. We studied the responses of two American maize inbreds, Mp708 and Tx601, to WCR infestation using biomechanical, molecular, biochemical analyses, and laser ablation tomography. Previous studies performed on several inbreds indicated that these two maize genotypes differed in resistance to pests including fall armyworm (Spodoptera frugiperda) and WCR. Our data confirmed that Mp708 shows resistance against WCR, and demonstrates that the resistance mechanism is based in a multi-trait phenotype that includes increased resistance to cutting in nodal roots, stable root growth during insect infestation, constitutive and induced expression of known herbivore-defense genes, including ribosomal inhibitor protein 2 (rip2), terpene synthase 23 (tps23) and maize insect resistance cysteine protease-1 (mir1), as well high constitutive levels of jasmonic acid and production of (E)-ß-caryophyllene. In contrast, Tx601 is susceptible to WCR. These findings will facilitate the use of Mp708 as a model to explore the wide variety of mechanisms and traits involved in plant defense responses and resistance to herbivory by insects with several different feeding habits.

Root anatomical phenes predict root penetration ability and biomechanical properties in maize (Zea Mays).

The ability of roots to penetrate hard soil is important for crop productivity but specific root phenes contributing to this ability are poorly understood. Root penetrability and biomechanical properties are likely to vary in the root system dependent on anatomical structure. No information is available to date on the influence of root anatomical phenes on root penetrability and biomechanics. Root penetration ability was evaluated using a wax layer system. Root tensile and bending strength were evaluated in plant roots grown in the greenhouse and in the field. Root anatomical phenes were found to be better predictors of root penetrability than root diameter per se and associated with smaller distal cortical region cell size. Smaller outer cortical region cells play an important role in stabilizing the root against ovalization and reducing the risk of local buckling and collapse during penetration, thereby increasing root penetration of hard layers. The use of stele diameter was found to be a better predictor of root tensile strength than root diameter. Cortical thickness, cortical cell count, cortical cell wall area and distal cortical cell size were stronger predictors of root bend strength than root diameter. Our results indicate that root anatomical phenes are important predictors for root penetrability of high-strength layers and root biomechanical properties.