Root hairs are a key trait for improving the acquisition of phosphorus (P) by plants. However, it is not known whether root hairs provide significant advantage for plant growth under combined soil stresses, particularly under conditions that are known to restrict root hair initiation or elongation (e.g. compacted or high-strength soils). To investigate this, the root growth and P uptake of root hair genotypes of barley, Hordeum vulgare L. (i.e. genotypes with and without root hairs), were assessed under combinations of P deficiency and high soil strength. Genotypes with root hairs were found to have an advantage for root penetration into high-strength layers relative to root hairless genotypes. In P-deficient soils, despite a 20% reduction in root hair length under high-strength conditions, genotypes with root hairs were also found to have an advantage for P uptake. However, in fertilized soils, root hairs conferred an advantage for P uptake in low-strength soil but not in high-strength soil. Improved root-soil contact, coupled with an increased supply of P to the root, may decrease the value of root hairs for P acquisition in high-strength, high-P soils. Nevertheless, this work demonstrates that root hairs are a valuable trait for plant growth and nutrient acquisition under combined soil stresses. Selecting plants with superior root hair traits is important for improving P uptake efficiency and hence the sustainability of agricultural systems.

Annual pasture legume species can vary more than 3-fold in their critical external phosphorus (P) requirement (i.e. P required for 90% of maximum yield). In this work we investigated the link between root morphology, P acquisition and critical external P requirement among pasture species. The root morphology acclimation of five annual pasture legumes and one grass species to low soil P availability was assessed in a controlled-environment study. The critical external P requirement of the species was low (Dactylis glomerata L., Ornithopus compressus L., Ornithopus sativus Brot.), intermediate (Biserrula pelecinus L., Trifolium hirtum All.) or high (Trifolium subterraneum L.). Root hair cylinder volumes (a function of root length, root hair length and average root diameter) were estimated in order to assess soil exploration and its impact on P uptake. Most species increased soil exploration in response to rates of P supply near or below their critical external P requirement. The legumes differed in how they achieved their maximum root hair cylinder volume. The main variables were high root length density, long root hairs and/or high specific root length. However, total P uptake per unit surface area of the root hair cylinder was similar for all species at rates of P supply below critical P. Species that maximised soil exploration by root morphology acclimation were able to prolong access to P in moderately P-deficient soil. However, among the species studied, it was those with an intrinsic capacity for a high root-hair-cylinder surface area (i.e. long roots and long root hairs) that achieved the lowest critical P requirement.

The effect of soil acidity on root and rhizosheath development in wheat and barley seedlings was investigated in an acid Ferrosol soil to which various amounts of lime (CaCO 3 ) were applied to modify soil Al concentrations (pH (CaCl 2 ): 4.22 to 5.35 and Al (CaCl 2 extract): 17.7 to 0.4 mg kg −1 soil; respectively), and Ferrosol soil from an adjacent location at the same site which had a higher Al concentration (pH 4.19; 29.2 mg kg −1 Al). The cereal lines were selected on the basis of differences in their rate of root growth, Al-resistance and root hair morphology. Root morphology was assessed after 7 days of growth. The length of fine (mainly lateral) roots of Al-sensitive genotypes was more sensitive to soil Al concentrations than that of the coarse (mainly primary) roots. The experiments demonstrated that even where root growth was protected by expression of the TaALMT1 gene for Al-resistance, root-soil contact was diminished by soil acidity because root hair length (in many lines), and root hair density and rhizosheath formation (all lines) were adversely affected by soil acidity. In the case of Al-sensitive lines, fine root growth and rhizosheath mass were reduced over much the same range of soil Al concentrations (i.e. >3-6 mg kg −1 Al). Although Al-resistant lines could maintain fine root length under these conditions, they were similarly unable to maintain rhizosheath mass. This finding may help to explain why Al-resistant wheats which yield relatively well in deep acid soils, may also benefit from application of lime to the surface layers of the soil.

Selecting plants with improved root hair growth is a key strategy for improving phosphorus-uptake efficiency in agriculture. While significant inter- and intra-specific variation is reported for root hair length, it is not known whether these phenotypic differences are exhibited under conditions that are known to affect root hair elongation. This work investigates the effect of soil strength, soil water content (SWC) and soil particle size (SPS) on the root hair length of different root hair genotypes of barley. The root hair and rhizosheath development of five root hair genotypes of barley (Hordeum vulgare L.) was compared in soils with penetrometer resistances ranging from 0.03 to 4.45 MPa (dry bulk densities 1.2-1.7 g cm(-3)). A "short" (SRH) and "long" root hair (LRH) genotype was selected to further investigate whether differentiation of these genotypes was related to SWC or SPS when grown in washed graded sand. In low-strength soil (<1.43 MPa), root hairs of the LRH genotype were on average 25 % longer than that of the SRH genotype. In high-strength soil, root hair length of the LRH genotype was shorter than that in low-strength soil and did not differ from that of the SRH genotype. Root hairs were shorter in wetter soils or soils with smaller particles, and again SRH and LRH did not differ in hair length. Longer root hairs were generally, but not always, associated with larger rhizosheaths, suggesting that mucilage adhesion was also important. The root hair growth of barley was found to be highly responsive to soil properties and this impacted on the expression of phenotypic differences in root hair length. While root hairs are an important trait for phosphorus acquisition in dense soils, the results highlight the importance of selecting multiple and potentially robust root traits to improve resource acquisition in agricultural systems.

Perennial pasture species are important for sustainable pasture systems; yet some species display poor persistence on acid soils. This work investigated the effect of soil acidity on primary root length and root-hair and rhizosheath development of five perennial grass genotypes varying in acid-soil resistance. Plants were grown in three low-P acid soils that were limed (CaCO 3 ) to modify soil pH (0.01 M CaCl 2 extractable) from <4.3 to >5.0 and Al 3+ concentrations from ≥ 16 to <4 mg kg −1 (0.01 M CaCl 2 extractable). Root morphology of tall wheatgrass (Thinopyrum ponticum (Podp.) Z.-W. Liu & R.-C Wang), phalaris (Phalaris aquatica L.), cocksfoot (Dactylis glomerata L.) and weeping grass (Microlaena stipoides Labill. R. Br.) was assessed after 20-25 days growth. The root length of tall wheatgrass and phalaris cv Sirosa was sensitive to acidity, with lateral root length more sensitive to acidity than seminal root length. Lime increased the root-hair length and density of both acid-soil sensitive and resistant genotypes (cocksfoot and an acidresistant line of phalaris) and root-hair length was positively correlated to an increase in rhizosheath size per unit root length. Restricted root length and poor root-soil contact of acid-soil sensitive genotypes may explain their low yield and relatively poor persistence on acid soils. The improvement to root-hair and rhizosheath development of genotypes that are acidsoil resistant in terms of root length demonstrates the benefit of using resistant genotypes in conjunction with liming to manage acid soils. Weeping grass was exceptional in its ability to maintain root length, roothair and rhizosheath development in acid soil.

Molecular techniques present a new opportunity to study roots and their interactions in soil. Extraction and quantification of species-specific DNA directly from soil allows direct identification of roots in mixed swards reducing the need for labour-intensive methods to recover and identify individual roots. DNA was extracted directly from up to 0.5 kg of soil and the presence of individual species quantified using speciesspecific probes with quantitative real-time PCR. A range of plant and soil factors influenced the DNA content measured in roots and it was necessary to account for these influences when converting DNA amount to root mass. The utility of the method for quantitative root studies was demonstrated in an experiment to investigate the effect of lime on root growth of acid-soil resistant and sensitive perennial grasses grown together in an aluminium-toxic soil. The root mass of an acid-soil resistant species was unaffected by lime application, whereas that of an acid-soil sensitive species was restricted by soil acidity. Molecular techniques present a promising tool for quantification of root mass directly in soil and have applications for field studies involving mixed species of plants.

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Background and Aims: This work aimed to quantify the critical external requirement for phosphorus (P) (i.e. external P concentration required for 90% of maximum yield) for a number of temperate legume species and understand differences in dry matter allocation, P distribution and P acquisition efficiency among these species. Methods: Shoot and root growth of five legume and one grass species was assessed in response to six rates of P mixed into the top 45 mm of soil in a pot experiment. Dactylis glomerata and Trifolium subterraneum were used as benchmark species; they are commonly grown together in mixed temperate pastures and have low and high critical external requirements for P, respectively. Growth was compared with four potential alternative legume species: Ornithopus compressus, Ornithopus sativus, Biserrula pelecinus and Trifolium hirtum, that have root morphologies better suited to soil exploration and nutrient acquisition than that of Trifolium subterraneum. Results: Dactylis glomerata, Ornithopus compressus and Ornithopus sativus had maximum yields equal to or greater than Trifolium subterraneum but achieved this at rates of P less than half that of Trifolium subterraneum. Biserrula pelecinus and Trifolium hirtum had critical P requirements between that of Trifolium subterraneum and the Ornithopus species, but also had lower yields. Root dry matter of Dactylis glomerata and the Ornithopus species in the fertilised soil layer was only marginally changed in response to low P supply. In contrast, Trifolium subterraneum, Trifolium hirtum and to a lesser extent Biserrula pelecinus markedly increased root dry matter allocation to this soil layer. Species with lower critical P requirements were able to take up more P per unit root dry mass than those with higher critical P requirements, particularly at lower levels of P addition. Conclusions: The high P acquisition efficiencies of the Ornithopus species and Dactylis glomerata were likely to have contributed to their low critical external P requirements. It was surmised that differences in root morphology traits underpin the differences in acclimation to low P stress and P acquisition efficiency among the species.