Rebecca E. Haling

Root hairs improve root penetration, root–soil contact, and phosphorus acquisition in soils of different strength

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.

Effect of soil acidity, soil strength and macropores on root growth and morphology of perennial grass species differing in acid‐soil resistance

It is unclear whether roots of acid-soil resistant plants have significant advantages, compared with acid-soil sensitive genotypes, when growing in high-strength, acid soils or in acid soils where macropores may allow the effects of soil acidity and strength to be avoided. The responses of root growth and morphology to soil acidity, soil strength and macropores by seedlings of five perennial grass genotypes differing in acid-soil resistance were determined, and the interaction of soil acidity and strength for growth and morphology of roots was investigated. Soil acidity and strength altered root length and architecture, root hair development, and deformed the root tip, especially in acid-soil sensitive genotypes. Root length was restricted to some extent by soil acidity in all genotypes, but the adverse impact of soil acidity on root growth by acid-soil resistant genotypes was greater at high levels of soil strength. Roots reacted to soil acidity when growing in macropores, but elongation through high-strength soil was improved. Soil strength can confound the effect of acidity on root growth, with the sensitivity of acid-resistant genotypes being greater in high-strength soils. This highlights the need to select for genotypes that resist both acidity and high soil strength.

Root morphological traits that determine phosphorus-acquisition efficiency and critical external phosphorus requirement in pasture species

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.

Field application of a DNA-based assay to the measurement of roots of perennial grasses

Background and aimsDNA-based methods present new opportunities for overcoming the difficulties of accurately identifying and quantifying roots of different plant species in field soils. In order to quantify species-specific root biomass from measurements of DNA, consideration needs to be given to replication and ability to recover roots for calibration purposes in order to account for spatial, temporal and inter- and intra-species variation in DNA content of roots and distribution of roots within the soil profile.MethodsThis paper develops the field application of a DNA-based technique for direct quantification of roots in soils. The method was applied to a field experiment to investigate differences in root growth of acid-soil resistant and sensitive genotypes of perennial pasture grasses in an acid soil. DNA was extracted directly from soil and species-specific DNA was quantified using quantitative real-time PCR prior to estimation of root biomass.ResultsRoot growth of the perennial grasses was quantified using the DNA-based technique, although separate calibration procedures were needed to convert DNA content to root mass for each species, soil layer and sampling date. Compared to acid-soil resistant genotypes, lesser root growth in acid soil layers and reduced above-ground dry matter production was observed for acid-soil sensitive genotypes.ConclusionsThe DNA-based method allowed genotypic differences in root growth to be assessed directly in soil and was advantageous for rapid processing of a large number of samples. However, high replication was still required to overcome spatial variability and separate calibrations were required for different species and soil depths across sampling times. The technique demonstrated greater root growth of acid-soil resistant perennial grasses which was beneficial for their establishment and persistence.

Ecological Succession, Hydrology and Carbon Acquisition of Biological Soil Crusts Measured at the Micro-Scale

The hydrological characteristics of biological soil crusts (BSCs) are not well understood. In particular the relationship between runoff and BSC surfaces at relatively large (>1 m2) scales is ambiguous. Further, there is a dearth of information on small scale (mm to cm) hydrological characterization of crust types which severely limits any interpretation of trends at larger scales. Site differences and broad classifications of BSCs as one soil surface type rather than into functional form exacerbate the problem. This study examines, for the first time, some hydrological characteristics and related surface variables of a range of crust types at one site and at a small scale (sub mm to mm). X-ray tomography and fine scale hydrological measurements were made on intact BSCs, followed by C and C isotopic analyses. A ‘hump’ shaped relationship was found between the successional stage/sensitivity to physical disturbance classification of BSCs and their hydrophobicity, and a similar but ‘inverse hump’ relationship exists with hydraulic conductivity. Several bivariate relationships were found between hydrological variables. Hydraulic conductivity and hydrophobicity of BSCs were closely related but this association was confounded by crust type. The surface coverage of crust and the microporosity 0.5 mm below the crust surface were closely associated irrespective of crust type. The δ 13C signatures of the BSCs were also related to hydraulic conductivity, suggesting that the hydrological characteristics of BSCs alter the chemical processes of their immediate surroundings via the physiological response (C acquisition) of the crust itself. These small scale results illustrate the wide range of hydrological properties associated with BSCs, and suggest associations between the ecological successional stage/functional form of BSCs and their ecohydrological role that needs further examination.

Spatial variability in pH and key soil nutrients: is this an opportunity to increase fertiliser and lime-use efficiency in grazing systems?

Abstract. Nutrient-use efficiency is a key issue for grazing systems in Australia. Spatial variability in soil pH and nutrients at the sub-paddock scale may affect the efficiency of utilisation of, and provide an opportunity for, site-specific management (SSM) of fertiliser and soil ameliorants. However, there has been little research exploring the potential for SSM in grazing systems. This study examines the spatial variability of soil test pH, phosphorus (P), potassium (K) and sulfur (S) in two typical pasture fields (a native and an improved) on the Northern Tablelands of New South Wales and evaluates the potential for SSM based on a comparison with critical values. In both fields, the overall paddock mean from a grid survey containing >80 samples for pH, P, K and S (0–10 cm) exceeded the critical values, suggesting that the addition of fertiliser or lime was not required. However, considerable sub-paddock-scale variability was observed, with CV ranging from 35% to 66% for the key nutrients (P, K and S). The Sprengel–Liebig Law of the Minimum was applied to evaluate the proportion of each field constrained by one or more soil characteristics. Up to 55% of the improved paddock and 78% of the native pasture was potentially responsive to amendments. The results of this study suggest that SSM of fertilisers and ameliorants could provide substantial improvements in productivity and possibly reductions in fertiliser use. The development and application of appropriate systems and tools to effectively quantify this spatial variability remain a challenge, coupled with management strategies that optimise the placement of amendments and account for the variability in other production limiting factors.

Effect of competition from a C4 grass on the phosphorus response of a subtropical legume

Abstract. Tropical pasture systems are typically dominated by C4 grasses growing on nitrogen (N) deficient soils. Under these conditions, N2-fixing legumes should have a competitive advantage, yet low legume contents are often reported in these systems. This work investigates whether below-ground competition for phosphorus (P) is limiting the ability of legumes to compete in swards of C4 grasses when grown in a sand matrix. The external P requirement of a subtropical legume (butterfly pea, Clitoria ternatea L.) and a C4 grass (buffel grass, Cenchrus ciliaris L.) were initially determined in a P-response experiment. Four rates of P (4.6–78.2 mg P kg–1 of Colwell P) were subsequently selected to investigate the growth response of the butterfly pea when grown with and without competition from a sward of N-deficient buffel grass. Shoot dry matter was determined over successive cuts and P uptake determined at the final harvest at 72 days. Buffel grass dominated the mixed swards and reduced the shoot dry matter production of the butterfly pea by >50% relative to the pure swards. A significant difference in the soil P response curve and shoot P uptake of butterfly pea was not detected between pure swards and those with competition from buffel grass. The ability of C4 grasses to acquire and convert resources (i.e. light, water and nutrients) more efficiently into shoot dry matter is likely to be a major factor resulting in grass-dominated pastures in tropical systems.

Differences in nutrient foraging among Trifolium subterraneum cultivars deliver improved P-acquisition efficiency

Background and aimsThe growth and root morphology responses to soil phosphorus (P) fertility by five cultivars of Trifolium subterraneum (a temperate annual pasture legume) were examined to assess whether differences in root morphology and/or root acclimation to P stress influenced P-acquisition by the clover varieties, or their critical P requirements (i.e. the rate of P supply needed for maximum shoot yield).MethodsThe clovers were grown as microswards in soil with P stratified in a topsoil layer to mimic growth conditions and soil P availability in a pasture. Yield and P content of shoots, and roots from the topsoil and subsoil layers was determined after 5 weeks growth in a controlled-environment cabinet. The lengths, diameters, and root hair lengths of nutrient foraging roots from the topsoil layer were quantified.ResultsThe shoot yield of the cultivars was similar when grown with high soil P fertility. However, the cultivars varied up to 1.5-fold in their ability to yield at low levels of soil P supply, and by 1.6-fold in their critical P requirements. All cultivars acclimated to low P soil by increasing root length density in the topsoil but those that yielded relatively well did so by maintaining a large root hair cylinder volume (i.e. they explored more soil) under low soil P conditions. This was associated with maintenance of dry matter allocations to topsoil roots and higher specific root lengths. Both factors assisted development of high root length density for nutrient foraging. Root acclimation responses to P were compared among the cultivars at equivalent relative shoot yields to assess the influence of plant P stress as a trigger for nutrient foraging. The least P-efficient cultivars slowed their allocation of dry matter to foraging roots at lower levels of P stress.ConclusionsThe results suggest variation within T. subterraneum for root proliferation and specific root length could be targeted in breeding for improved P-acquisition efficiency.

The carboxylate composition of rhizosheath and root exudates from twelve species of grassland and crop legumes with special reference to the occurrence of citramalate

AimsLow-molecular-weight organic anions (carboxylates) influence rhizosphere processes and may enhance plant phosphorus acquisition. We examined the root exudate profile of a range of pasture and grain legumes and focused on the little-investigated carboxylate, citramalate.MethodsTwelve species of pasture legumes and herbs, including four Lotus spp. and two crop legumes, Cicer arietinum and Lupinus albus, were grown in a glasshouse for six weeks. The composition and amounts of carboxylates were measured in rhizosheath soil as well as in root exudates from roots washed free of their rhizosheath.ResultsCitrate and malate were found in the rhizosheath of all species. However, citramalate was present in the rhizosheath of only Lotus species (10–82% of total exuded carboxylates). Cicer arietinum had the largest amount of carboxylates in its rhizosheath and fastest rate of carboxylate exudation into the trap solution.ConclusionsCitrate and malate were found in the rhizosheath of all species in this study, but citramalate was only found in the root exudates and rhizosheath of Lotus spp. Further investigation into the role of citramalate in Lotus spp. is merited.

Intrinsic capacity for nutrient foraging predicts critical external phosphorus requirement of 12 pasture legumes

Abstract. The mainstream pasture legume species such as Trifolium subterraneum, T. repens and annual Medicago spp. used in the temperate pasture systems of southern Australia have high critical external requirements for phosphorus (P) (i.e. P required to achieve 90% of maximum yield). This work aimed to identify alternative pasture legume species that could be used in systems with lower P input. Shoot and root biomass of 12 species of pasture legume was measured in response to seven rates of P applied to the top 48 mm of soil in a pot experiment. Most species had maximum yields similar to T. subterraneum, but some required only one-third of the applied P to achieve this. The critical external P requirement of the species, ranked from lowest to highest, was as follows: Ornithopus compressus = O. sativus < Biserrula pelecinus < T. michelianum = T. vesiculosum = T. glanduliferum < T. hirtum = Medicago truncatula = T. purpureum = T. incarnatum < T. spumosum = T. subterraneum. An ability to maximise soil exploration through a combination of high root-length density, high specific root length and long root hairs (i.e. a large specific root-hair-cylinder volume) was associated with a low critical external P requirement. The results indicate that Ornithopus spp. could be used to achieve productive, low P-input pasture systems.