Effects of plant pathogens on population dynamics and community composition in grassland ecosystems: two case studies

Grassland ecosystems comprise a major portion of the earth’s terrestrial surface, ranging from high-input cultivated monocultures or simple species mixtures to relatively unmanaged but dynamic systems. Plant pathogens are a component of these systems with their impact dependent on many interacting factors, including grassland species population dynamics and community composition, the topics covered in this paper. Plant pathogens are affected by these interactions and also act reciprocally by modifying their nature. We review these features of disease in grasslands and then introduce the 150-year long-term Park Grass Experiment (PGE) at Rothamsted Research in the UK. We then consider in detail two plant-pathogen systems present in the PGE, Tragopogon pratensis-Puccinia hysterium and Holcus lanata-Puccinia coronata. These two systems have very different life history characteristics: the first, a biennial member of the Asteraceae infected by its host-specific, systemic rust; the second, a perennial grass infected by a host-non-specific rust. We illustrate how observational, experimental and modelling studies can contribute to a better understanding of population dynamics, competitive interactions and evolutionary outcomes. With Tragopogon pratensis-Puccinia hysterium, characterised as an “outbreak” species in the PGE, we show that pathogen-induced mortality is unlikely to be involved in host population regulation; and that the presence of even a short-lived seed-bank can affect the qualitative outcomes of the host-pathogen dynamics. With Holcus lanata-Puccinia coronata, we show how nutrient conditions can affect adaptation in terms of host defence mechanisms, and that co-existence of competing species affected by a common generalist pathogen is unlikely.

The inclusion of forage mixtures in the diet of growing dairy heifers: impacts on digestion, energy utilisation, and methane emissions

Intensive farming focusing on monoculture grass species to maximise forage production has led to a reduction in the extent and diversity of species-rich grasslands. However, plant communities with higher species number (richness) are a potential strategy for more sustainable production and mitigation of greenhouse gas (GHG) emissions. Research has indicated the need to understand opportunities that forage mixtures can offer sustainable ruminant production systems. The objective of the two experiments reported here were to evaluate multiple species forage mixtures in comparison to ryegrass-dominant pasture, when conserved or grazed, on digestion, energy utilisation, N excretion, and methane emissions by growing 10–15 month old heifers. Experiment 1 was a 4 × 4 Latin square design with five week periods. Four forage treatments of: (1) ryegrass (control); permanent pasture with perennial ryegrass (Lolium perenne); (2) clover; a ryegrass:red clover (Trifolium pratense) mixture; (3) trefoil; a ryegrass:birdsfoot trefoil (Lotus corniculatus) mixture; and (4) flowers; a ryegrass:wild flower mixture of predominately sorrel (Rumex acetosa), ox-eye daisy (Leucanthemum vulgare), yarrow (Achillea millefolium), knapweed (Centaurea nigra) and ribwort plantain (Plantago lanceolata), were fed as haylages to four dairy heifers. Measurements included digestibility, N excretion, and energy utilisation (including methane emissions measured in respiration chambers). Experiment 2 used 12 different dairy heifers grazing three of the same forage treatments used to make haylage in experiment 1 (ryegrass, clover and flowers) and methane emissions were estimated using the sulphur hexafluoride (SF6) tracer technique. Distribution of ryegrass to other species (dry matter (DM) basis) was approximately 70:30 (clover), 80:20 (trefoil), and 40:60 (flowers) for experiment 1. During the first and second grazing rotations (respectively) in experiment 2, perennial ryegrass accounted for 95 and 98% of DM in ryegrass, and 84 and 52% of DM in clover, with red clover accounting for almost all of the remainder. In the flowers mixture, perennial ryegrass was 52% of the DM in the first grazing rotation and only 30% in the second, with a variety of other flower species occupying the remainder. Across both experiments, compared to the forage mixtures (clover, trefoil and flowers), ryegrass had a higher crude protein (CP) content (P < 0.001, 187 vs. 115 g kg −1 DM) and DM intake (P < 0.05, 9.0 vs. 8.1 kg day −1). Heifers in experiment 1 fed ryegrass, compared to the forage mixtures, had greater total tract digestibility (g kg −1) of DM (DMD; P < 0.008, 713 vs. 641) and CP (CPD, P < 0.001, 699 vs. 475), and used more intake energy (%) for body tissue deposition (P < 0.05, 2.6 vs. −4.9). For both experiments, heifers fed flowers differed the most compared to the ryegrass control for a number of measurements. Compared to ryegrass, flowers had 40% lower CP content (P < 0.001, 113 vs. 187 g kg −1), 18% lower DMD (P < 0.01, 585 vs. 713 g kg −1), 42% lower CPD (P < 0.001, 407 vs. 699 g kg −1), and 10% lower methane yield (P < 0.05, 22.6 vs. 25.1 g kg −1 DM intake). This study has shown inclusion of flowers in forage mixtures resulted in a lower CP concentration, digestibility and intake. These differences were due in part to sward management and maturity at harvest. Further research is needed to determine how best to exploit the potential environmental benefits of forage mixtures in sustainable ruminant production systems.

Digestion, rumen fermentation and circulating concentrations of insulin, growth hormone and IGF-1 in steers fed diets based on different proportions of maize silage and grass silage

Replacing grass silage with maize silage results in a fundamental change in the ratio of structural to non-structural carbohydrates with commensurate changes in rumen fermentation patterns and nutrient utilisation. This study investigated the effects of feeding four forage mixtures, namely grass silage (G); 67 g/100 g grass silage133 g/100 g maize silage (GGM); 67 g/100 g maize silage133/100 g grass silage (MMG); maize silage (M) to four ruminally and duodenally canulated Holstein Friesian steers. All diets were formulated to be isonitrogenous (22.4 g N/kg DM) using a concentrate mixture. Dietary dry matter (DM) and organic matter (OM) digestibility increased with ascending maize silage inclusion (P,0.1) whereas starch and neutral detergent fibre digestibility declined (P,0.05). Ratio of non-glucogenic to glucogenic precursors in the rumen fluid increased with maize silage inclusion (P,0.01) with a commensurate reduction in rumen pH (P,0.05). Mean circulating concentrations of insulin were greatest and similar in diets MMG and GGM, lower in diet M and lowest in diet G (P,0.01). There were no effects of diet on the mean circulating concentration of growth hormone (GH), or the frequency, amplitude and duration of GH pulses, or the mean circulating concentrations of IGF-1. Increasing levels of DM, OM and starch intakes with the substitution of grass silage with maize silage affected overall digestion, nutrient partitioning and subsequent circulating concentrations of insulin.

Apparent digestibility and nitrogen utilisation of diets based on maize and grass silage fed to beef steers

Substituting grass silage with maize silage in forage mixtures may result in one forage influencing the nutritive value of another in terms of whole tract nutrient digestibility and N utilisation. This experiment investigated effects of four forage combinations being, grass silage (G); 67 g/100 g grass silage + 33 g/100 g maize silage (GGM); 67 g/100 g maize silage + 33 g/100 g grass silage (MMG); maize silage (M). All diets were formulated to be isonitrogenous (22.4 g N/kg dry matter [DM]) using a concentrate mixture. Ration digestibility and N balance was determined using 7 Holstein Friesian steers (mean body weight 411.0 +/- 120.9 kg) in a cross-over design. Inclusion of maize silage in the diet had a positive linear effect on forage and total DM intake (P = 0.001), and on apparent DM and organic matter digestibility (both P = 0.048). Regardless of the silage ratio used, the metabolisable energy concentration of maize silage was calculated to be higher than that of grass silage (P = 0.058), and linearly related to the relative proportions of the two silages in the forage mixture. Inclusion of maize silage in the diet resulted in a linear decline in the apparent digestibility of starch (P = 0.022), neutral detergent fibre (P < 0.001) and acid detergent fibre (P = 0.003). Nitrogen retention, expressed as amount retained per day or in terms of body weight (g/100 kg) increased linearly with maize inclusion (P = 0.047 and 0.046, respectively). Replacing grass silage with maize silage caused linear responses according to the proportions of each forage in the diet, and that there were no associative effects of combining forages. (C) 2004 Elsevier B.V. All rights reserved.

A field experiment to recreate species rich hay meadows using regional seed mixtures

Species rich semi-natural grasslands are an important but threatened habitat throughout Europe and much of the former area has been lost since the 1950s. However, in some countries large areas have been preserved and the demand for meadow recreation by sowing seed mixtures is increasing. In the White Carpathians Protected Landscape Area (Czech Republic) the use of commercial seed mixtures is undesirable and the use of regional mixtures has been investigated. The costs for seeding large areas are high and lower cost techniques are needed. In 1999 a field experiment was set up to investigate the establishment of hay meadow vegetation comparing sowing a regional mixture all over a plot with sowing narrow 2.5 In strips of regional seed mixtures into a matrix of a commercial grass mixture or into natural regeneration. The results after five seasons showed good establishment of the sown species in the meadow treatment. Spread of sown species from the sown strips into the surrounding matrix occurred but the cover of species was lower in the commercial grass matrix compared with the natural regeneration matrix. Colonisation of some plots by unsown desirable grassland species from adjacent grassland habitats also occurred, but more species colonised the natural regeneration matrix than the commercial grasses or the sown meadow matrix itself. Overall, the results indicate that, in appropriate situations, sown strips can provide a lower cost but slower and longer-term alternative to field scale sowing of regional seed mixtures for recreation of hay meadow vegetation. (C) 2007 Elsevier Ltd. All rights reserved.

Investigating the phytotoxicity of the graminicide fluazifop-P-butyl against native UK wildflower species

BACKGROUND: The selective graminicide fluazifop-P-butyl is used for the control of grass weeds in dicotyledonous crops, and commonly applied in amenity areas to reduce grass productivity and promote wildflower establishment. However, evidence suggests that fluazifop-P-butylmight also have phytotoxic effects on somenon-target plants. This study investigates the effects of fluazifop-P-butyl on the emergence, phytotoxicity and above-ground biomass of nine perennial wildflower species and two grass species, following pre- and post-emergent applications at half, full and double label rates in a series of glasshouse experiments. RESULTS: While pre- and post-emergent applications of fluazifop-P-butyl caused reductions in seedling emergence and increased phytotoxicity on native wildflower and grass species, these effects were temporary for the majority of wildflower species tested, and generally only occurred at the double application rate. No differences in biomass were observed at any of the rates, suggesting good selectivity and no long-term effects of fluazifop-P-butyl application on the wildflower species from either pre-emergent or post-emergent applications. CONCLUSION: These results have direct relevance to the management of amenity areas for biodiversity, as they confirm the suitability of these wildflower species for inclusion in seed mixtures where fluazifop-P-butyl is to be applied to control grass productivity.

The grass-free lawn: management and species choice for optimum ground cover and plant diversity

In Britain, managed grass lawns provide the most traditional and widespread of garden and landscape practices in use today. Grass lawns are coming under increasing challenge as they tend to support a low level of biodiversity and can require substantial additional inputs to maintain. Here we apply a novel approach to the traditional monocultural lawnscape by replacing grasses entirely with clonal perennial forbs. We monitored changes in plant coverage and species composition over a two year period and here we report the results of a study comparing plant origin native, non-native and mixed) and mowing regime. This allows us to assess the viability of this construct as an alternative to traditional grass lawns. Grass-free lawns provided a similar level of plant cover to grass lawns. Both the mowing regime and the combination of species used affected this outcome, with native plant species seen to have the highest survival rates, and mowing at 4cm to produce the greatest amount of ground coverage and plant species diversity within grass-free lawns. Grass-free lawns required over 50% less mowing than a traditionally managed grass lawn. Observations suggest that plant forms that exhibited: a) a relatively fast growth rate, b) a relatively large individual leaf area, and c) an average leaf height substantially above the cut to be applied, were unsuitable for use in grass-free lawns. With an equivalent level of ground coverage to grass lawns, increased plant diversity and a reduced need for mowing, the grass-free lawn can be seen as a species diverse, lower input and potentially highly ornamental alternative to the traditional lawn format.

The influence of plant species number on productivity, ground coverage and floral performance in grass-free lawns

The grass-free lawn is a novel development in modern ornamental horticulture where the traditional monoculture of grass is replaced by a variety of mowing-tolerant clonal forbs. It brings floral aesthetics and a diverse species approach to the use of lawn space. How the number of constituent forb species affects the aesthetic and structural performance of grass-free lawns was investigated using grass-free lawns composed of four, six and twelve British native clonal perennial forb species. Lawn productivity was seen to increase with increasing species number but the relationship was not linear. Plant cover was dynamic in all lawn types, varied between years and was not representative of individual species' floral performance. The behaviour of component species common to all lawns suggested that lawns with 12 species show greater structural stability than the lawns with a lower species number. Visual performance in lawns with the greatest species number was lower than in lawns with fewer species, with increasing variety in floral size and individual species floral productivity leading to a trade-off between diversity and floral performance. Individual species were seen to have different aesthetic functions in grass-free lawns either by providing flowers, ground coverage or both.

The grass-free lawn: floral performance and management implications

Grass lawns are a ubiquitous feature of urban green-space throughout much of the temperate world. Species poor and intensively managed, lawns are ecologically impoverished, however environmentally aware lawn owners are reluctant to implement alternatives due to aesthetic concerns. Developing an alternative lawn format which is both biodiversity friendly and aesthetically pleasing is an imperative for urban greening. We suggest that such an alternative can be provided by replacing the grass lawn by a forb-based mix. To advance this, we tested the floral performance of three groups of clonal perennial forbs (native, non-native and mixed), each maintained using standard lawn management mowing regimes. Our findings show that both the frequency of mowing and the height at which mowing is applied influence floral performance and lawn aesthetics. Species origin was found to influence floral productivity, floral visibility and floral variety within grass-free lawns, with native species providing the greatest floral performance. The behaviour and management of grass lawns was not found to be a suitable analogue for the management of grass-free lawns and grass-free lawns are sufficiently different from grass lawns to require an entirely original management approach. We suggest that the grass-free lawn can provide an aesthetically and environmentally relevant replacement for the ubiquitous and ecologically-poor grass lawn.

Factors affecting distribution and mobility of trace elements (Cu, Pb, Zn) in a perennial grapevine (Vitis vinifera L.) in the Champagne region of France

Soil and Vitis vinifera L (coarse and fine roots, leaves, berries) concentration and geochemical partitioning of Cu, Pb and Zn were determined in a contaminated calcareous Champagne plot to assess their mobility and transfer. Accumulation ratios in roots remained low (0.1-0.4 for Cu and Zn, <0.05 for Pb). Differences between elements resulted from vegetation uptake strategy and soil partitioning. Copper, significantly associated with the oxidisable fraction (27.8%), and Zn with the acid soluble fraction (33.3%), could be mobilised by rhizosphere acidification and oxidisation, unlike Pb, essentially contained in the reducible fraction (72.4%). Roots should not be considered as a whole since the more reactive fine roots showed higher accumulation ratios than coarse ones. More sensitive response of fine roots, lack of correlation between chemical extraction results and vegetation concentrations, and very limited translocation to aerial parts showed that fine root concentrations should be used when assessing bioavailability. (C) 2008 Elsevier Ltd. All rights reserved.

Response of organically managed grassland to available phosphorus and potassium in the soil and supplementary fertilization: field trials using grass-clover leys cut for silage

Effective use and recycling of manures together with occasional and judicious use of supplementary fertilizing materials forms the basis for management of phosphorus (P) and potassium (K) within organic farming systems. Replicated field trials were established at three sites across the UK to compare the supply of P and K to grass-clover swards cut for silage from a range of fertilizing materials, and to assess the usefulness of routine soil tests for P and K in organic farming systems. None of the fertilizing materials (farmyard manure, rock phosphate, Kali vinasse, volcanic tuff) significantly increased silage yields, nor was P offtake increased. However, farmyard manure and Kali vinasse proved effective sources of K to grass and clover in the short to medium term. Available P (measured as Olsen-P) showed no clear relationship with crop P offtake in these trials. In contrast, available K (measured by ammonium nitrate extraction) proved a useful measurement to predict K availability to crops and support K management decisions.

Host plant recognition by the root feeding clover weevil, Sitona lepidus (Coleoptera : Curculionidae)

This study investigated the ability of neonatal larvae of the root-feeding weevil, Sitona lepidus Gyllenhal, to locate white clover Trifolium repens L. (Fabaceae) roots growing in soil and to distinguish them from the roots of other species of clover and a co-occurring grass species. Choice experiments used a combination of invasive techniques and the novel technique of high resolution X-ray microtomography to non-invasively track larval movement in the soil towards plant roots. Burrowing distances towards roots of different plant species were also examined. Newly hatched S. lepidus recognized T. repens roots and moved preferentially towards them when given a choice of roots of subterranean clover, Trifolium subterraneum L. (Fabaceae), strawberry clover Trifolium fragiferum L. (Fabaceae), or perennial ryegrass Lolium perenne L. (Poaceae). Larvae recognized T. repens roots, whether released in groups of five or singly, when released 25 mm (meso-scale recognition) or 60 mm (macro-scale recognition) away from plant roots. There was no statistically significant difference in movement rates of larvae.

MEASUREMENT AND MODELING OF THE TOXICITY OF BINARY MIXTURES IN THE NEMATODE CAENORHABDITIS ELEGANS-A TEST OF INDEPENDENT ACTION

Ecological risk assessments must increasingly consider the effects of chemical mixtures on the environment as anthropogenic pollution continues to grow in complexity. Yet testing every possible mixture combination is impractical and unfeasible; thus, there is an urgent need for models that can accurately predict mixture toxicity from single-compound data. Currently, two models are frequently used to predict mixture toxicity from single-compound data: Concentration addition and independent action (IA). The accuracy of the predictions generated by these models is currently debated and needs to be resolved before their use in risk assessments can be fully justified. The present study addresses this issue by determining whether the IA model adequately described the toxicity of binary mixtures of five pesticides and other environmental contaminants (cadmium, chlorpyrifos, diuron, nickel, and prochloraz) each with dissimilar modes of action on the reproduction of the nematode Caenorhabditis elegans. In three out of 10 cases, the IA model failed to describe mixture toxicity adequately with significant or antagonism being observed. In a further three cases, there was an indication of synergy, antagonism, and effect-level-dependent deviations, respectively, but these were not statistically significant. The extent of the significant deviations that were found varied, but all were such that the predicted percentage effect seen on reproductive output would have been wrong by 18 to 35% (i.e., the effect concentration expected to cause a 50% effect led to an 85% effect). The presence of such a high number and variety of deviations has important implications for the use of existing mixture toxicity models for risk assessments, especially where all or part of the deviation is synergistic.

Intake, growth and meat quality of steers given diets based on varying proportions of maize silage and grass silage

Simmental × Holstein-Friesian steers were offered four forage diets. These comprised grass silage (G); proportionately 0·67 grass silage, proportionately 0·33 maize silage (GGM); 0·33 grass silage, 0·67 maize silage ( MMG); maize silage ( M) from 424 (s.d. = 11·5) kg to slaughter at a minimum weight of 560 kg. Forages were mixed and offered ad libitum. Steers were offered 2 kg of a concentrate daily, the concentrate being formulated such that all steers had similar crude protein intakes across dietary treatments. A sample of steers was slaughtered at the beginning of the experimental period to allow the calculation of the rate of gain of the carcass and its components. Carcass dissection of a sample of steers allowed the development of a prediction equation of carcass composition based on thoracic limb dissection of all carcasses. Forage dry matter intake and live-weight gain increased linearly as maize silage replaced grass silage in the forage mixture, resulting in improvements in food conversion ratio (all P = 0·001). Killing-out proportion increased with maize silage inclusion ( P < 0·001) but fat and conformation scores did not differ significantly between diets. However, increasing maize inclusion in the diet resulted in a greater weight ( P = 0·05) and proportion ( P = 0·008) of fat in the carcass, and significant increases in internal fat deposition. The inclusion of maize led to a progressive increase in the daily gains of carcass ( P < 0·001), and significant increases in the daily gains of both fat ( P < 0·001) and lean tissue ( P < 0·001). Fat colour was more yellow in cattle given diets G and GGM than diets MMG and M ( P < 0·001) and colour intensity was lower on diet M than the other three diets ( P < 0·001). There were no significant differences in any aspects of eating quality between diets. Therefore, maize silage has the potential to reduce the time taken for finishing beef animals to achieve slaughter weight with no apparent detrimental effects on subsequent meat quality.

Soil moisture mediates association between the winter North Atlantic Oscillation and summer growth in the Park Grass Experiment

Several aspects of terrestrial ecosystems are known to be associated with the North Atlantic Oscillation (NAO) through effects of the NAO on winter climate, but recently the winter NAO has also been shown to be correlated with the following summer climate, including drought. Since drought is a major factor determining grassland primary productivity, the hypothesis was tested that the winter NAO is associated with summer herbage growth through soil moisture availability, using data from the Park Grass Experiment at Rothamsted, UK between 1960 and 1999. The herbage growth rate, mean daily rainfall, mean daily potential evapotranspiration (PE) and the mean and maximum potential soil moisture deficit (PSMD) were calculated between the two annual cuts in early summer and autumn for the unlimed, unfertilized plots. Mean and maximum PSMD were more highly correlated than rainfall or PE with herbage growth rate. Regression analysis showed that the natural logarithm of the herbage growth rate approximately halved for a 250 mm increase in maximum PSMD over the range 50-485 mm. The maximum PSMD was moderately correlated with the preceding winter NAO, with a positive winter NAO index associated with greater maximum PSMD. A positive winter NAO index was also associated with low herbage growth rate, accounting for 22% of the interannual variation in the growth rate. It was concluded that the association between the winter NAO and summer herbage growth rate is mediated by the PSMD in summer.