Model predicting dynamics of biomass, structure and digestibility of herbage in managed permanent pastures. 1. Model description

To investigate seasonal and annual interactions between management and grassland dynamics, a simple mechanistic model of the dynamics of production, structure and digestibility in permanent pastures was constructed. The model is designed to respond to various defoliation regimes, perform multiple‐year simulations and produce simple outputs that are easy to use as inputs for a model of ruminant livestock production. Grassland communities are described using a set of average functional traits of their constituent grass groups. The sward is subdivided into four structural compartments: green leaves and sheath, dead leaves and sheath, green stems and flowers, and dead stems and flowers. Each compartment is characterized by its biomass, age and digestibility. Only above‐ground growth is modelled, using a light‐utilization efficiency approach modulated by a seasonal pattern of storage and mobilization of reserves. Ageing of plant parts is driven by cumulative thermal time from 1 January and by biomass flows. Age affects senescence, abscission and digestibility of green compartments and, therefore, the quality of green leaves and stems can increase or decrease over time in relation to net growth and defoliation dynamics. The functional traits having the greatest impact on model outputs are seasonal effects, period of reproductive growth and effects of temperature on photosynthetic efficiency. The functional traits of the grass groups were parameterized for temperate pastures of the Auvergne region in France. The other model inputs are few: proportion of functional groups, basic weather data (incident photosynthetically active radiation, mean daily temperature, precipitation and potential evapotranspiration) and site characteristics (nitrogen nutrition index, soil water‐holding capacity). In the context of a whole‐farm simulator, the model can be applied at a field scale.     

The indirect estimation of the digestibility in cattle of herbage from Belgian permanent pasture

Relationships between herbage organic matter digestibility measured in vivo , and faecal nitrogen concentration were determined. Measurements of in vivo digestibility were made on twenty-nine herbages, cut from mixed improved permanent pastures with Lolium perenne as the predominant grass and offered ad libitum to cattle. The faecal nitrogen index predicted the herbage digestibility with good accuracy, and the equations obtained should have application for such determinations on Belgian pastures.     

Animal production evaluation of herbage varieties. 1. Comparison of Aurora with Frances, Talbot and Melle perennial ryegrasses when grown alone and with clover

Comparative herbage characteristics and sheep production (ewes and lambs until weaning and lambs thereafter) from swards of Aurora (very early flowering), Frances (early flowering), Talbot (intermediate flowering) and Melle (late flowering) varieties of perennial ryegrass ( Lolium perenne ) were studied under a continuous variable stocking management based on sward surface height guidelines. The varieties were assessed as grass-only (215 kg N ha⁻¹ fertilizer) and grass/clover (83 kg N ha⁻¹ fertilizer) pastures.
Over 2 years (1985-86) total annual lamb production per hectare from grass-only swards of Aurora was 19% more than that from Frances despite similar herbage productivity. Differences between the varieties in lamb output were more pronounced as grass/clover swards, with Aurora producing 29 and 18% more lamb than Melle and Talbot respectively and with Frances also giving 16% more than Melle. Overall lamb production from grass/clover swards was 10% more than that from the grass-only pastures, mainly due to 69% advantage in individual lamb growth rates after weaning. Herbage organic matter digestibility, during the post-weaning period, was higher on grass/clover than on grass-only swards, and similarly with Aurora versus the other varieties. Under a frequent cut simulated grazing regime the relative herbage productivity of the four varieties differed, with Aurora 13% less productive than Frances.
The results are discussed in relation to the limitations of assessing performance under cutting, and the significance of the interaction in animal productivity between the two sward types.     

Seasonal dynamics of biomass production and herbage quality of three grasslands with contrasting functional compositions

Classifying grasses into plant functional types (PFT) according to their strategy of nutrient capture (PFT A, B and b) or nutrient conservation (PFT C) is claimed to predict variability in biomass production and herbage quality in grasslands. PFT A, B and b are characterized by higher production than PFT C, with higher nutritive value, which decreases rapidly. PFT C is characterized by lower nutritive value, which remains more stable in time. The aim of this study was to quantify the consequences of PFT composition at the community level on the seasonal dynamics of biomass and herbage quality for livestock feed. We studied three grasslands located in the same area with contrasting PFT composition and classified PFT A, PFT Bb and PFT C. The dynamics of vegetation stages, dry‐matter yield and nutritive value were followed over three vegetation cycles in 2007–2008. In the first cycle, biomass accumulation was high for the PFT A and PFT Bb grasslands and the high quality of herbage declined rapidly with the reproductive development. By contrast, biomass accumulation was slow in the PFT C grassland, with a lower quality that declined slowly, consistent with the later reproductive development. For regrowth cycles, the same ranking of grasslands was observed for biomass accumulation, and differences in digestibility were similar to those observed at the beginning of the first vegetation cycle. Combining type of grassland with sum of temperature for the first cycle and regrowth days for the second and third vegetation cycles provided quantitative models of herbage biomass and quality.     

Response of permanent and reseeded grassland to fertilizer nitrogen. 1. Herbage production and herbage quality

The productivity of permanent swards of mixed species composition was compared with that of sown Lolium perenne , cv. Melle at five fertilizer-N rates (0, 150, 300, 450 and 900 kg N ha⁻¹) and with L. perenne/Trifolium repens , cv. Grasslands Huia at 0 kg N ha⁻¹. The investigation was conducted under two cutting frequencies at sixteen sites in England and Wales, representing a range of grassland environments.
Annual total herbage dry matter (DM) production from both permanent and reseeded swards increased with successive increments of fertilizer-N up to 450 kg N ha⁻¹. Herbage DM production from reseeded swards in the first year after sowing was signficantly higher than from the permanent swards, at all fertilizer-N treatments. In subsequent years the production advantage of the L. perenne reseeds was maintained only at the higher N rates, though sown L. perenne/T. repens was the most productive sward type at 0 N. Average differences in modified acid-detergent fibre suggested small advantages in herbage quality to the reseeded swards.
It is concluded that, while reseeded swards are more productive in the year after sowing, many permanent swards are capable of high levels of production and that reseeding to a L. perenne sward cannot always be justified, particularly for grassland receiving low or moderate inputs of fertilizer-N.     

The effects of level of Molinia utilization on diet selection and herbage intake by cattle grazing Molinia grassland

Groups of mature, non-lactating, cows grazed two Molinia -dominant grassland communities in central and southern Scotland during six consecutive summers. Two treatments, designed to use either 33% or 66% of the estimated annual Molinia leaf production by grazing to different leaf lengths, were imposed at each site. Grazing was restricted to the period of Molinia growth each season. During the first 4 years, diet composition, diet digestibility and herbage organic matter intake were determined during either one or two measurement periods each year. There were differences between sites in the floristic content of the sward and these differences were reflected in the diet selected by the cattle. Cows grazing the taller (33% utilization) plots had higher percentages of Molinia , grass stem, sheath and inflorescences and lower percentages of broad-leaved grasses, sedges, rushes and dead herbage in the diet than those grazing the shorter (66% utilization) plots. Differences between the floristic composition of the sward and the diet were explicable by (a) the height at which cattle grazed in relation to the distribution of components within the sward or (b) the selective grazing of small areas dominated by a particular species. The organic matter digestibility of diets differed between sites but there was no significant difference in digestibility or organic matter intake between the treatments. On average less than 50 d grazing was provided by the experimental sites each year. During this period the liveweight gains of cows grazing the two treatments did not differ significantly. The implications of these results for the management of Molinia -dominant communities are discussed.     

Defoliation and productivity of a phalaris-subterranean clover sward, and the influence of grazing experience on sheep intake

Swards of Phalaris aquatica-Trifolium subterraneum were subjected to four defoliation treatments—zero, low (11 sheep ha⁻¹) and high (22 sheep ha⁻¹) stocking rates, and weekly cutting. At high stocking rate the annual grass Hordeum leporinum dominated while clover was dominant at low and zero stocking rates. Weekly cutting suppressed species other than clover and so failed to simulate grazing.
There were similarities in net herbage production between zero and lightly grazed swards and between heavily grazed and repeatedly cut swards. Net herbage production decreased in the order undisturbed sward < lightly grazed sward < heavily grazed sward < repeatedly cut sward.
When sheep grazed swards where herbage mass was low their daily consumption of herbage, and therefore liveweight change, depended on their recent grazing experience. Sheep accustomed to swards where herbage mass was low ate more because they grazed for much longer each day than unaccustomed sheep, although they selected a diet of similar digestibility.     

Seasonal productivity and nutritive value of temperate grasses found in semi-natural pastures in Europe: responses to cutting frequency and N supply

Monocultures of thirteen perennial C₃ grass species that co‐occur in temperate semi‐natural grassland communities in Europe were compared in a factorial field design of two levels of N supply and two levels of cutting frequency. Above‐ground yield of dry matter (DM), crude protein (CP) concentration and pepsin‐cellulase DM digestibility of herbage were measured in two successive years. Species was the largest source of variability in yield of DM and DM digestibility of herbage, while CP concentration of herbage responded more to management factors. The highest mean DM yields and values of DM digestibility of herbage were achieved in spring for Festuca arundinacea and in autumn for Phleum pratense. Poa trivialis and Festuca rubra had the lowest DM yield and DM digestibility values, respectively, regardless of seasons and treatments. For all species CP concentration in herbage increased in response to an increase in cutting frequency and N supply by an average of 46 and 34 g kg^?1 DM respectively. Differences between years and seasons indicated the importance of plant phenology on nutritional variables and the influence of environmental factors on species performance. Species ranking was compared according to their annual digestible DM and CP yields. The results show that some grasses have a nutritive value which is comparable to that of forages selected for high yields.     

Defoliation patterns and their implications for the management of vegetative tropical pastures to control intake and diet quality by cattle

This study assessed the use of pasture attributes to control daily intake and diet quality during progressive defoliation on pastures of Axonopus catarinensis. Three consecutive 12‐day grazing treatments of progressive defoliation were conducted with Brahman cross‐steers. Daily forage intake and defoliation dynamics were assessed using a pasture‐based method. The treatments differed in initial sward height (33, 44 and 61 cm) and herbage mass (1030, 1740 and 2240 kg ha^?1). The post‐grazing residual sward height, at which forage intake decreased, appeared to increase with the initial sward height (12·3, 14·6 and 15·5 cm). Steers grazed up to four distinctive grazing strata in all treatments. The depth and herbage mass content of the top grazing stratum were at least five times higher than the lower grazing strata in all treatments. This explains why forage intake decreased when the top grazing stratum was removed in approximately 93% of the pasture area in all treatments, equivalent to approximately 7% of the pasture area remaining ungrazed. We conclude that the residual ungrazed area of the pasture, rather than residual sward height, can be used to develop grazing management strategies to control forage intake and diet quality in a wide range of pasture conditions.     

THE ENERGY RETENTION OF LAMBS GRAZING S37 COCKSFOOT, S23 RYEGRASS AND S24 RYEGRASS AT LOW GRAZING PRESSURE

A grazing experiment to measure the feeding values of 4-week-old regrowths of S37 cocksfoot, S23 ryegrass and S24 ryegrass is described. A low grazing pressure was maintained on the pastures to ensure that animal production was not limited by herbage availability. Each sward was grazed by a group of 12 lambs under "worm-free" conditions. Increases in empty body weight, dry body weight and energy were measured by the comparative slaughter technique.
The liveweight gain of the cocksfoot-fed lambs was the same as the ryegrass-fed groups during the first half of the experiment but was lower in August and September. The final liveweight gains on S23 and S24 ryegrass were 14% higher than on S37 cocksfoot, but in terms of energy retention S23 ryegrass was 18% and S24 7% better than S37 cocksfoot. Digestibility measurements of herbage cut from the plots being grazed showed a positive correlation with energy retention by the lambs. Energy retention was negatively correlated with the proportion of acetic acid in the rumen adds, a fraction which was itself correlated with the soluble carbohydrate content of the herbage, but not with the digestibility of the herbage.
When interpreting animal-production results, intake, digestibility and rumen volatilc-fativ-acid proportions, should be taken into account. Because of the inadequacy of present methods of estimating the herbage intake of grazing animals, measurement of body-energy gains is not justified and empty-liveweight gains are sufficiently precise to measure relative energy retentions at pasture.     

The effects of pesticide application on upland permanent pasture

An estimate of the level of pest damage on enclosed upland permanent pasture in England and Wales was made by reducing invertebrate populations with insecticides at thirteen well-distributed sites and measuring the effect on herbage growth over a 3-year period. Potential pests were fewer in number than on lowland pastures and there was a wide range in herbage yield between sites. Insecticide treatment significantly increased annual yields at only three of the sites, and significantly decreased yield at two of the sites. Insect damage to grassland appears to be much less important in upland than in lowland areas: this may be due to the relative scarcity of frit-fly in these areas, but this could change if its preferred host, ryegrass, were encouraged by intensification of sward management in the uplands. At all but one of the sites insecticide treatment led to considerable soil compaction, probably as a consequence of suppressing earthworm populations.     

Effect of herbage mass and allowance on sward characteristics,milk production,intake and rumen volatile fatty acid concentration

The objective of this study was to examine the effect of herbage mass and daily herbage allowance (DHA) on sward characteristics and animal performance, dry‐matter intake, rumen pH and volatile fatty acid production of unsupplemented spring‐calving dairy cows throughout the main grazing season. Sixty‐eight Holstein‐Friesian dairy cows were randomly assigned across four treatments (n = 17) in a 2 × 2 factorial design. Two swards were created with different levels of pre‐grazing herbage mass [allocated above 4 cm (>4 cm); 1700 kg DM ha^?1 (medium; M) or 2200 kg DM ha^?1 (high; H)] and two levels of DHA (>4 cm; 16 or 20 kg DM per cow d^?1). An additional eight lactating ruminally cannulated Holstein–Friesian dairy cows were randomly assigned to each treatment in a replicated 4 × 4 Latin square design. Sward and animal measurements were collected across four periods each of 1 week duration in April and May (PI) and July and August (PII). Maintaining the medium‐mass sward across the season improved the nutritive value of the sward in the latter part of the grazing season compared with high‐mass swards, thus resulting in increased animal intakes and milk production throughout PII. The higher organic matter digestibility of the medium‐compared with high‐masses during PII indicates that grazing severity and herbage mass in the spring to mid‐summer period will determine sward quality parameters in the late summer period.     

Changes in the area of permanent grassland and its implications for the provision of bioenergy: Slovakia as a case study

Throughout Europe, grasslands are managed primarily for agricultural production but also provide a range of ecosystem services, the magnitude of which is influenced by their area, management and the abiotic properties of the ecosystem. The grassland area in Europe has been affected by significant changes in recent decades, including abandonment and conversion to arable land. This study presents an assessment of changes in the permanent grassland area with reference to Slovakia and provides an assessment of the bioenergy potential of permanent grassland surplus to agricultural needs. In 2015, of the total permanent grassland area (868,000 ha) 42% had ceased to be used as feed for milk or meat production and had been abandoned. Results from field experiments show that low‐level improvement of permanent grasslands, e.g., by oversowing commonly sown agricultural grassland species and low‐dose fertilization (up to 30 kg N ha^?1 year^?1) could provide herbage biomass of 1.7 million tonnes of herbage dry matter per year. This would be sufficient to generate an energy output of approximately 31.3–31.9 PJ/year (4.5% of the current gross inland energy consumption of Slovakia). Low‐intensity use of the currently surplus permanent grassland is also considered to have potential beneficial outcomes in terms of delivery of ecosystem services. Results are also discussed in the context of countries beyond the case‐study area.     

Effects of livestock breed and grazing intensity on biodiversity and production in grazing systems. 1. Nutritive value of herbage and livestock performance

Reduction of grazing intensity and the use of traditional instead of commercial breeds has frequently been recommended to meet biodiversity and production goals in sustainable grazing systems in Europe. To test the impact of such practices across a range of contrasting grassland types, integrated measurements of foraging behaviour, agronomic production and botanical, structural and invertebrate biodiversity were made over three years on four sites in the UK, Germany, France and Italy. The sites in the UK and Germany were mesotrophic grassland with high productivity and low to moderate initial levels of plant diversity, and were grazed by cattle. The French site was a semi‐natural, species‐rich grassland grazed by cattle. The Italian site contained a wider range in plant diversity, from species‐rich to mesotrophic grassland, and was grazed by sheep. The treatments were: MC, moderate grazing intensity with a commercial breed – this was designed to utilize herbage growth for optimum livestock production; LC, lenient grazing intensity with a commercial breed – this was designed to increase biodiversity by not fully utilizing herbage growth; and LT, lenient grazing intensity with a traditional breed – this was also designed to increase biodiversity. Neither fertilizers nor pesticides were applied. The nutritive value of the herbage and the performance of the livestock were measured. Mean stocking rates were proportionately 0·30–0·40 lower and mean sward heights and herbage mass on offer were 0·30–0·50 higher on the LC and LT treatments compared with the MC treatment. The proportion of live and dead material, and leaves and stems in the herbage, its chemical composition and nutritive value were little affected by the treatments. Individual livestock performance, measured as liveweight gain, showed no consistent response to treatment. In Germany, performance on the MC treatment was slightly lower than on the LC and LT treatments but no such difference was found on the sites in the other countries. Livestock breed did not have a strong effect on livestock performance. In the UK and France the traditional breeds had a lower performance but this was not the case in Germany or Italy. Livestock performance per ha of the LC and LT treatments was up to 0·40 lower than of the MC treatment. It is concluded that biodiversity‐targeted extensive grazing systems have potential to be integrated into intensive livestock production systems because the individual livestock performance reaches a similar level compared to a moderate grazing intensity. Traditional breeds did not have a production advantage over commercial breeds on extensively managed pastures.     

Interactions between leaf lifespan and defoliation frequency in temperate and tropical pastures: a review

In continuously stocked swards or pastures the frequency at which individual tillers and individual leaves are defoliated by ruminant livestock, relative to leaf lifespan of the grass species within the sward, determines the proportion of each leaf defoliated before senescence, and hence the efficiency of harvesting of herbage. In this paper, sets of data obtained in a range of climatic conditions and with a range of grass species are used in order to document this relationship. It is shown that the frequency of defoliation of individual tillers or individual leaves is closely linked to the average stocking density used within a period of time for maintaining a steady state sward or pasture height, herbage mass or leaf area index. Consequently, any decrease in herbage growth rate should lead to a decrease in the efficiency of harvesting of herbage and then to a more than proportional decrease in total herbage consumption by ruminant livestock. These effects will be more important for grass species having short leaf lifespan than for species with long lifespan. In rotational stocking, the link between herbage growth rate and frequency of defoliation of leaves can be broken by controlling the grazing interval, so any decrease in herbage growth would not be systematically associated with a decrease in efficiency of harvesting of herbage. Rotational stocking should be more efficient than continuous stocking in low herbage production conditions, while in high herbage production systems rotational and continuous stocking would have similar efficiency. The implications of these conclusions for the management of swards and pastures to meet different objectives are discussed briefly.     

Variations in herbage mass and digestibility, and the grazing behaviour and herbage intake of adult cattle and weaned calves

Two experiments are described in which groups of lactating cows, non-lactating cattle and weaned calves grazed a sequence of swards varying in maturity and herbage mass under strip grazing management at a daily herbage allowance of 60 g dry matter per kg live weight. Lactating cows ate 43% and 76% more herbage than non-lactating cattle of similar weight in the two experiments but herbage OM intakes per unit live weight by the calves and lactating cows were similar.
Variations in diet digestibility and herbage intake in the lactating and non-lactating cattle with changes in sward conditions were similar. In the first experiment the calves were experienced grazers; the variation in the digestibility of the herbage selected was less in the calves than the adult cattle, but the variation in herbage intake was greater. The calves in the second experiment were younger, and they were inexperienced grazers; their ability to increase herbage intake in response to changing sward conditions was poorer than that of adult cattle, though variations in diet digestibility were similar in all classes.
The practical implications of these results are discussed briefly.     

Performance and environmental effects of forage production on sandy soils. III. Energy efficiency in forage production from grassland and maize for silage

Based on experimental data gathered in a research project on nitrogen fluxes in intensive dairy farming in Northern Germany, an analysis of fossil energy input and energy efficiency in forage production from permanent grassland and maize for silage was conducted. Field experiments comprised different defoliation systems and different rates of mineral N fertilizer and slurry application. Each change from grazing to cutting in grassland systems reduced the energy efficiency. Energy efficiency consistently decreased with increasing rates of mineral N application. In the production of maize for silage, maximum energy efficiency was obtained with an application of 50 kg N ha^?1 from slurry only. Net energy yields of maize for silage were much higher than that of grassland when compared at the same level of fossil energy and nitrogen fertilizer input. Considering both nitrate‐leaching losses and a necessary minimum quantity of grass herbage in a well‐balanced ration, it is suggested that a high proportion of maize for silage in combination with N‐unfertilized grass/clover swards used in a mixed cutting/grazing system represents a good trade‐off between the leaching of nitrates and energy efficiency.     

Scaling up of a mechanistic dynamic model in a GIS environment to model temperate grassland production at the regional scale

Regional‐scale ecological processes are mediated by processes occurring at local scales. Spatially explicit models are needed to understand the broad‐scale consequences of a large number of local processes, driven by factors which are heterogeneous at a broad scale. A Geographic Information System (GIS)‐based mechanistic model, which can be used as a flexible tool to investigate the regional‐scale effects of changes in environmental factors on herbage production from a Lolium perenne sward, was developed and used to investigate the consequences of aggregating driving variables at different resolutions. The model allows rigorous scaling up of previously existing field‐based modelling approaches within a GIS context. The model's mechanistic approach allows flexibility in the simulation of the separate effects of environmental driving variables and of cutting regimes. The driving variables (temperature, solar radiation, available soil moisture and soil nitrogen status) are scaled up using geostatistical techniques. The model is used to evaluate weekly changes in herbage production under environmental conditions, variable both in space and time, under different cutting regimes. It is shown that, in this model, aggregation at 1 km² is a good compromise between accuracy and practical feasibility, and that, while ignoring heterogeneity over many square kilometres can induce large errors, their magnitude and direction also depend on the model response curve to an input variable. The results obtained were consistent with the known trends in influential environmental factors. The programming within a GIS makes the model flexible in its application and, therefore, makes it easy to apply at a variety of scales.     

GrazeIn: a model of herbage intake and milk production for grazing dairy cows. 2. Prediction of intake under rotational and continuously stocked grazing management

Decision support tools to help dairy farmers gain confidence in grazing management need to be able to predict performance of grazing animals with easy‐to‐obtain variables on farm. This paper, the second of a series of three, describes the GrazeIn model predicting herbage intake for grazing dairy cows. The model of voluntary intake described in the first paper is adapted to grazing situations taking account of sward characteristics and grazing management, which can potentially affect intake compared to indoor feeding. Rotational and continuously stocked grazing systems are considered separately. Specific effects of grazing management on intake were quantified from an extensive literature review, including the effect of daily herbage allowance and pre‐grazing herbage mass in rotational grazing systems, sward surface height in continuously stocked grazing systems, and daily time at pasture in both grazing systems. The model, based on iterative procedures, estimates many interactions between cows, supplements, sward characteristics and grazing management. The sensitivity of the prediction of herbage intake to sward and management characteristics, as well as the robustness of the simulations and an external validation of the GrazeIn model with an independent data set, is presented in a third paper.