Richard A. Culvenor

Plant and microbial strategies to improve the phosphorus efficiency of agriculture

BackgroundAgricultural production is often limited by low phosphorus (P) availability. In developing countries, which have limited access to P fertiliser, there is a need to develop plants that are more efficient at low soil P. In fertilised and intensive systems, P-efficient plants are required to minimise inefficient use of P-inputs and to reduce potential for loss of P to the environment.ScopeThree strategies by which plants and microorganisms may improve P-use efficiency are outlined: (i) Root-foraging strategies that improve P acquisition by lowering the critical P requirement of plant growth and allowing agriculture to operate at lower levels of soil P; (ii) P-mining strategies to enhance the desorption, solubilisation or mineralisation of P from sparingly-available sources in soil using root exudates (organic anions, phosphatases), and (iii) improving internal P-utilisation efficiency through the use of plants that yield more per unit of P uptake.ConclusionsWe critically review evidence that more P-efficient plants can be developed by modifying root growth and architecture, through manipulation of root exudates or by managing plant-microbial associations such as arbuscular mycorrhizal fungi and microbial inoculants. Opportunities to develop P-efficient plants through breeding or genetic modification are described and issues that may limit success including potential trade-offs and trait interactions are discussed. Whilst demonstrable progress has been made by selecting plants for root morphological traits, the potential for manipulating root physiological traits or selecting plants for low internal P concentration has yet to be realised.

Strategies and agronomic interventions to improve the phosphorus-use efficiency of farming systems

Phosphorus (P)-deficiency is a significant challenge for agricultural productivity on many highly P-sorbing weathered and tropical soils throughout the world. On these soils it can be necessary to apply up to five-fold more P as fertiliser than is exported in products. Given the finite nature of global P resources, it is important that such inefficiencies be addressed. For low P-sorbing soils, P-efficient farming systems will also assist attempts to reduce pollution associated with P losses to the environment. P-balance inefficiency of farms is associated with loss of P in erosion, runoff or leaching, uneven dispersal of animal excreta, and accumulation of P as sparingly-available phosphate and organic P in the soil. In many cases it is possible to minimise P losses in runoff or erosion. Uneven dispersal of P in excreta typically amounts to ~5% of P-fertiliser inputs. However, the rate of P accumulation in moderate to highly P-sorbing soils is a major contributor to inefficient P-fertiliser use. We discuss the causal edaphic, plant and microbial factors in the context of soil P management, P cycling and productivity goals of farms. Management interventions that can alter P-use efficiency are explored, including better targeted P-fertiliser use, organic amendments, removing other constraints to yield, zone management, use of plants with low critical-P requirements, and modified farming systems. Higher productivity in low-P soils, or lower P inputs in fertilised agricultural systems can be achieved by various interventions, but it is also critically important to understand the agroecology of plant P nutrition within farming systems for improvements in P-use efficiency to be realised.

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.

Persistence of winter-active phalaris breeding populations, cultivars and other temperate grasses in diverse environments of south-eastern Australia

Three winter-active populations of phalaris (Phalaris aquatica L.), selected over two generations for improved persistence under grazing, were evaluated with commercial cultivars of phalaris and other temperate perennial grasses from 1999–2003 in three environments of south-eastern Australia as part of a program to develop a cultivar for more sustainable pastures and to assess genotype × environment interaction. Grazed sites were located at Bulart in western Victoria, and Rye Park on the Southern Tablelands and Tamworth on the North West Slopes of New South Wales. At the conclusion of the experiment, the frequency of live plant base was highest at Rye Park despite soil acidity and drought. Significant variance among half-sib families in each population was also observed most frequently at this site. Frequency was intermediate at Bulart but lower than expected considering high soil fertility, probably because of high grazing pressure. Frequency was lowest at Tamworth where severe drought occurred from 2001 onwards. There was significant genotype × environment interaction for frequency among half-sib families. Significant common family variance for frequency across the Bulart and Rye Park sites was demonstrated, but not between Tamworth and either of the other sites in later years. The relationship between winter herbage mass potential and persistence differed with population and site, and was negative for one population at Bulart but positive for another population at Tamworth. Mean persistence of all families was 30% higher than winter-active controls at Rye Park and at least 40% higher at Bulart. Phalaris generally persisted better than cultivars of tall fescue (Festuca arundinacea Schreb.), cocksfoot (Dactylis glomerata L.) and perennial ryegrass (Lolium perenne L.) with some exceptions, particularly at Bulart. Development of a winter-active phalaris cultivar with improved persistence under grazing was considered possible for the Southern Tablelands and western Victorian environments with these populations but a separate program using additional germplasm will be needed for the North West Slopes environment.

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.

The effect of improved aluminium tolerance on establishment of the perennial grass, phalaris, on strongly acid soils in the field and its relation to seasonal rainfall

We examined the hypothesis that the higher aluminium (Al) tolerance of recent cultivars of the perennial grass, phalaris (Phalaris aquatica L.), improved establishment on strongly acid soils and that expression of this tolerance depended on seasonal rainfall conditions. Establishment under a range of conditions by a new cultivar (Advanced AT) with the highest Al tolerance yet achieved was compared with another relatively tolerant cultivar (Landmaster) and less tolerant cultivars and some alternative species on strongly acidic soils by sowing twice per year at six sites over 3 years (2004–06). Seedlings were considered established if they survived the first summer as shown by frequency and yield in the year after sowing. Low autumn rainfall in all years hindered weed control and delayed sowing. Under adequate rainfall in 2004 and 2005 sowings, Advanced AT and Landmaster established similarly and were denser and more productive, on average, than Holdfast and Sirosa, which in turn were higher than Australian II, an order related to Al tolerance measured in solution. Persistence in later years was ranked similarly. Under very adverse spring drought conditions in 2006, Advanced AT established better than Landmaster at the most acidic sites as hypothesised. Soil pH of sites where Advanced AT established better than Landmaster in 2006–07 was <4.2 in CaCl2. The results suggested that current recommendations on the tolerance of phalaris to acid soils are too conservative and that phalaris is more suitable for acid soils in southern Australia than these recommendations indicate.

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.

Persistence traits in perennial pasture grasses: the case of phalaris (Phalaris aquatica L.)

Abstract. Persistence is consistently claimed by Australian farmers as a high priority for perennial grasses in long-term pastures. Phalaris (Phalaris aquatica L.) is a productive perennial grass with proven persistence in south-eastern Australia. Nevertheless, factors that determine the persistence of pasture species in southern Australia related to climate (drought), soil (acidity), grazing pressure, and, importantly, their interaction can reduce persistence of phalaris and other species in various situations. These factors and their interactions are discussed in this review, and strategies to improve persistence with emphasis on plant breeding approaches are considered, with the most durable outcomes achieved when breeding and management options are employed concurrently. Two examples of breeding to improve persistence traits in phalaris are described. A program to improve acid-soil tolerance resulted first in the release of cv. Landmaster, and recently Advanced AT, which is the most aluminium (Al)-tolerant cultivar of phalaris to date. It was bred by recurrent selection on acid soils in a population containing genes from a related, more Al-tolerant species, P. arundinacea. The higher Al tolerance of cv. Advanced AT is of most benefit in more assured establishment on acid soils under variable moisture conditions and confers improved flexibility of sowing date. Cultivar Holdfast GT was bred to address complaints of poor persistence under heavy grazing by cultivars of the highly productive, winter-active type, since high grazing tolerance is needed to achieve profitable returns from developed pastureland. Evidence of good persistence under grazing for cv. Holdfast GT and possible tradeoffs with productivity are discussed. Maintaining high productivity under a predicted higher incidence of drought stress (climate change) and increasing areas of acid soils presents ongoing challenges for persistence in pastures.

Response to selection for grazing tolerance in winter-active populations of phalaris (Phalaris aquatica L.). 1. Persistence under grazing in three environments

Forage grass cultivars must have adequate grazing tolerance for use in the grazing systems for which they are intended. Response to 2 cycles of selection for persistence under heavy grazing pressure was examined in 3 winter-active breeding populations of the productive perennial grass, phalaris (Phalaris aquatica L.), from 2000 to 2003, at Bulart in western Victoria and Rye Park on the Southern Tablelands and Tamworth on the North-West Slopes of New South Wales. There was one continuously grazed and one rotationally grazed set of plots at Bulart to examine the effect of grazing management. All sites were affected by drought in later years but drought stress was most severe at Tamworth. A strongly positive linear response to selection was observed in an analysis of persistence measured as frequency of phalaris plant base across the 3 sites (excluding the rotational treatment at Bulart), but response interacted with site. By 2003, linear response averaged 14% frequency units or 34–40% proportional response per cycle at Rye Park and the continuously grazed plots at Bulart, sites that were environmentally suited to survival of phalaris. In contrast, response to selection was absent or slightly negative at Tamworth where conditions were drier and hotter. Populations responded similarly in analyses across all 3 sites but 1 population was less responsive when analyses were restricted to Bulart and Rye Park. Herbage mass measurements in 2001 at Bulart and 2002 at Rye Park indicated positive responses to selection, which were increasingly linked to frequency over time. Rotationally grazed plots at Bulart displayed higher frequency than continuously grazed plots after 3 years of grazing. The experiment showed that grazing tolerance was a heritable trait in the populations tested when environmental constraints were not limiting and that useful improvements in persistence had been obtained compared with existing cultivars.

Response to selection for grazing tolerance in winter-active populations of phalaris (Phalaris aquatica L.). 2. Correlated response in yield potential, plant characteristics, and alkaloid levels

Phenotypic changes in populations of perennial grasses are known to occur in response to natural or deliberate selection under grazing. These changes may have agronomic significance. Associated changes in morphology and yield potential of young stands in response to 2 cycles of selection for grazing tolerance were examined in 3 winter-active breeding populations of the perennial grass, phalaris (Phalaris aquatica L.). Levels of alkaloids which potentially could affect palatability were also examined. There was a decline in seedling growth and autumn and winter yield in spaced plants of 6.3–7.5% per cycle pooled across populations. Seedling growth measured in sown swards at 2 sites was not affected by selection. Visually estimated sward yield in the second year, ignoring large gaps, agreed with the spaced plant results but the decline was not significant at P ≤ 0.05 when herbage yield was measured by mowing, probably due to effects of plant density. On balance, it was concluded that a decline in individual plant yield of 6–7% per cycle had occurred but this could be compensated by higher density, particularly over time as differences in persistence under grazing developed. The most pronounced morphological response to selection under grazing was towards a more densely tillered growth habit, although one exception occurred. There was also a tendency towards a more prostrate growth habit and later heading, but this was significant only for the most erect and earliest maturing population. Two cycles of selection did not significantly affect summer activity or area of plant base in any population. Tryptamine alkaloids were below the level likely to affect palatability but tended to increase with selection, particularly in a very low tryptamine base population, which suggested that they may play a role in persistence. Because of potentially deleterious effects on yield, care is required in using this selection method. A balance of yield potential and grazing tolerance appropriate to the management system is needed.