Relationships between morphological and chemical characteristics of perennial ryegrass varieties and intake by sheep under continuous stocking management

Plots of five intermediate‐heading varieties of perennial ryegrass (Lolium perenne L.) [AberDove, Belramo and Glen (diploid); Twins (tetraploid); and AberExcel (tetraploid hybrid)] were continuously stocked with sheep to maintain a target sward surface height of 40–50 mm. Daily dry matter (DM) intake was significantly different (F‐value = 0·032) between the varieties, with the tetraploid hybrid AberExcel having the highest values for daily DM intake and intake rate during eating. Amongst the diploid varieties, intake rate tended to be higher for sheep grazing Glen. The varieties comprised a wide range in potential growth habit, from the relatively prostrate, highly tillered Glen to the more‐erect AberExcel and there were differences between them in the vertical distribution of leaves within the sward canopy. The leaves of AberExcel weighed 3·6 mg DM cm^?2 leaf area in contrast to the other varieties (4·3–5·3 mg DM cm^?2 leaf area) resulting in a high leaf area index (LAI) in relation to the green leaf mass. Intake rate was not significantly correlated with extended tiller and sheath tube lengths, partition of herbage mass, number of tillers per square metre or LAI. However, canonical variates analysis showed that there were significant differences between the varieties for the morphological and chemical factors examined. Other factors also need to be explored to explain these differences in ingestive behaviour in order to identify plant traits that are correlated with herbage intake rate. These are needed for varieties destined for grazing use, both during the breeding programme and their subsequent evaluation.     

Effects of perennial ryegrass (Lolium perenne L.) cultivars on herbage production, nutritional quality and herbage intake of grazing dairy cows

Four perennial ryegrass (Lolium perenne L.) cultivars were compared for differences in herbage production, nutritive value and herbage intake of dry matter (DM) during the summers of 2002 and 2003. Two paddocks were sown with pure stands of four cultivars in a randomized block design with three replicates. Each plot was subdivided into fourteen subplots (22 m × 6 m) which were grazed by one cow during 24 h. Twelve lactating dairy cows were assigned to one cultivar for a period of 2 weeks in a 4 × 4 Latin square experimental design; the experiment lasted 8 weeks in each year. Sward structure (sward surface height, DM yield, green leaf mass, bulk density and tiller density) and morphological characteristics were measured. The ash, neutral‐detergent fibre, acid‐detergent lignin, crude protein and water‐soluble carbohydrate concentrations, and in vitro digestibility of the herbage were measured. The sward was also examined for infestation by crown rust (Puccinia coronata f. sp. lolii). Herbage intake of dairy cows was estimated using the n‐alkane technique. Cultivar differences for all sward structural characteristics were found except for bulk density and tiller density in 2003. Cultivars differed for proportions of pseudostem, stem (in 2003 only) and dead material. The chemical composition of the herbage was different among cultivars, with the water‐soluble carbohydrate concentration showing large variation (>0·35). Cultivars differed in susceptibility to crown rust. Herbage intake differed among cultivars in 2002 (>2 kg DM) but not in 2003. Herbage intake was positively associated with sward height, DM yield and green leaf mass. Canopy morphology did not affect herbage intake. Crown rust affected herbage intake negatively. It was concluded that options for breeders to select for higher intake were limited. High‐yielding cultivars and cultivars highly resistant to crown rust were positively related with a high herbage intake.     

The effect of sward structure as influenced by ryegrass genotype on bite dimensions and short-term intake rate by dairy cows

Abstract The effects of genotypic variation in ryegrasses on sward structure, bite dimensions and intake rate by dairy cows were investigated. Two experiments were conducted. In Experiment 1, swards were in a vegetative state whereas, in Experiment 2, they were partly reproductive and were taller with higher herbage mass but lower leaf proportion than in Experiment 1. Applicability of relationships between sward structure and bite characteristics, previously established from artificial or hand‐constructed swards, to field conditions were tested. Additional short‐term intake rates and/or sward structural characteristics were considered as indicators of potential intake for use in protocols for the evaluation of grass varieties. Four cultivars were studied: AberElan, Twins (diploid and tetraploid perennial ryegrasses respectively), Polly, a hybrid ryegrass (perennial × Italian ryegrass) and Multimo (Italian ryegrass), each established in 200‐m² plots in four replicated blocks. Herbage intake rate was determined by short‐term liveweight change (taking account of insensible weight loss) using 16 dairy cows allocated to four balanced groups with each plot grazed by one group for a 1‐h assessment period. One block was grazed per day, over a 4‐d experimental period, with each group grazing each variety in a complete crossover design. Sward characteristics and bite rate were also measured in both experiments. Bite dimensions were subsequently estimated, with bite depth being determined as a function of extended tiller height (ETH) in both experiments. Within both experiments, bite mass and intake rate did not differ significantly between swards of different cultivars despite swards containing Multimo generally having a higher ETH and water‐soluble carbohydrate concentration and lower green leaf mass, sward bulk density and neutral‐detergent fibre concentration than the other swards. However, bite depth was significantly higher (P < 0·01) in swards containing Multimo swards than in the others and, in Experiment 1, bite depth, as a proportion of ETH, was higher in swards containing Multimo and lower in those containing Twins than in the other two cultivars, whereas there was no difference in Experiment 2. Taking both experiments together, the mean bite depth was 0·5 of ETH with sward bulk density accounting for almost half the variance in the relationship between bite depth and ETH. The bulk density of the bite (bite mass per unit bite volume), measured in Experiment 2, followed a similar pattern to sward bulk density, increasing in the order Multimo, Polly, AberElan and Twins. It is concluded that the relationships between sward characteristics and bite dimensions, derived from artificial swards, are applicable to field swards, although the range in natural ryegrass sward characteristics is usually not as wide as in experiments using artificial swards. Lack of precision in the measurement of short‐term intake and in sward‐based measurements is likely to preclude their use in the evaluation of grass varieties.     

Intake and behaviour responses by sheep to changes in sward characteristics under rotational grazing

A 10-week grazing experiment was conducted on a perennial ryegrass sward with lactating ewes and their twin lambs. Three paddocks were rotationally grazed with rest periods of from 4 to 5 weeks. Sward surface heights at the start of each grazing were 145, 259 and 250mni for treatments RG1, RG2 and RG3. A further four paddocks were maintained by continuous variable stocking (CS) at sward surface heights (SSHs) of about 30, 60, 90 and 120mm. Sward and animal measurements were made on the two different grazing managements as the RG swards were grazed down, giving measurements at similar sward heights for treatments RG and CS.
There was less green leaf and the total herbage mass present under RG was less than on CS swards at the same sward heights, demonstrating the differences in structure between rotationally and continuously grazed swards.
Regression analysis of animal factors on sward factors showed that grazing behaviour was more highly correlated with green leaf mass than SSH or any of the other sward measurements. On the RG swards, maximum intake per animal was reached at about 1500 kg green leaf mass ha⁻¹. A SSH of 60mm allowed the CS ewes to achieve the highest intake rate, but at this height the ewes on treatments RG2 and RG3 were restricted to approximately half this rate. The results suggest that green leaf mass or leaf area index, rather than sward surface height, could be used as a rational basis to relate intake of herbage to sward state for swards changing rapidly in leaf to stem ratio.     

A comparison of diploid and tetraploid perennial ryegrass and tetraploid ryegrass/white clover swards under continuous sheep stocking at controlled sward heights. 3. Sward characteristics and animal output, years 4–8

Results for years 4–8 of a long-term grazing experiment on swards of a diploid perennial ryegrass (Lolium perenne), var. Contender (D swards), a tetraploid ryegrass, var. Condesa (T swards) and Condesa with S184 white clover (Trifolium repens) (TC swards), direct sown in May 1987, are presented. The swards were continuously stocked with sheep from 1988 to 1990, as previously reported, and for a further 5 years, 1991–95, at a target sward surface height (SSH) of 4–6 cm. Control of sward height was successfully achieved by variable stocking, except in 1993 when paddocks were set stocked and the resulting mean SSH was 9·3 cm. Grass swards received on average 160 kg N ha^?1 year^?1; grass/clover swards were mainly not fertilized with N with the exception that they were given 30 kg N ha^?1 as a remedial mid-summer application during a period of low herbage mass on offer in 1994 and 1995. Mean white clover content of the swards fell from 18·2% of herbage dry-matter (DM) in 1992 to 8·5% in 1993, whereas stolon lengths fell from 120 to 58 m m^?2. A return to lower sward heights in 1994–95 resulted in an increase in white clover content to 12·8% by the final sampling in August 1995. Perennial ryegrass content of the grass swards remained high throughout (mean 96·7% in 1995). Perennial ryegrass tiller densities recorded in August 1991, 1993 and 1994 showed consistently significant (P < 0·001) sward differences (3-year mean 16 600, 13 700 and 10 100 perennial ryegrass tillers m^?2 for the D, T and TC swards). In 1994, the year after lax grazing, a low perennial ryegrass tiller density (9100 m^?2) and low white clover content (mean 4·3%) in the TC swards resulted in a much lower herbage bulk density than in the grass swards (April–July means 72, 94 and 44 kg OM ha^?1 cm^?1 for the D, T and TC swards). There was a consistent 40 g d^?1 increase in lamb liveweight gain on the TC swards over the T swards, except in 1994. In that year there was a reduction in lamb liveweight gain of 33 g d^?1 on the TC swards and a significant increase in ewe liveweight loss (117 g d^?1) associated with low herbage bulk density despite optimal sward height. Lamb output (kg liveweight ha^?1) on TC swards reflected white clover content, falling from a similar output to that produced from grass given 160 kg N ha^?1, at 18% white clover DM content, down to 60% of grass + N swards with around 5% clover. A 6% greater output from the T than the D swards was achieved mainly through higher stocking rate. The experiment demonstrated a rapid, loss in white clover under lax grazing, and showed that the relationship between performance and sward height is also dependent on herbage density. High lamb output from a grass/clover sward was only achieved when the clover content was maintained at 15–20% of the herbage DM.     

The effects of regrowth and maintenance height on a grass sward with a high density of tillers

A perennial ryegrass sward was grazed by sheep in April 1993 to a target sward surface height (SSH) of 3 cm to create a high density of grass tillers. From 3 May, the sheep were removed and small plots were established on the sward, when the average tiller density (± s.e.) was 35 900 ± 420 live tillers m⁻². Different regrowth treatments were then imposed by allowing plots to regrow to target SSHs of 6 cm (18 g dry matter (DM) M⁻²), 9 cm (78 g DM m⁻²), 12 cm (132 g DM m⁻²) or 15 cm (197 g DM m⁻²). The plots were then maintained by cutting at either 6 or 9 cm SSH until the end of the experiment on 30 September. Live tiller density was reduced by regrowth beyond 8.9 cm (78 g DM m⁻², P <0.001) and leaf-stem ratio and in vitro organic matter digestibility were reduced by regrowth beyond 6.1 cm (18 g DM m⁻², P < 0.05). The effect on live tiller density was sustained through the remainder of the season. From the beginning of June to the end of September, maintenance of SSH at an average of 9.1 cm compared with 6.4 cm also resulted in lower live tiller density, live-dead tiller ratio and leaf-stem ratio and higher herbage mass (at least P < 0. 05). There were significant interactions between regrowth SSH and maintenance SSH, so that leaf-stem ratio, live-dead tiller ratio and live tiller density were reduced by regrowth to a SSH of 16–4 cm followed by maintenance at 9.1 cm, compared with regrowth to 6.1 cm and maintenance at 6.4 cm.     

Defoliation pattern, foraging behaviour and diet selection by lactating dairy cows in response to sward height and herbage allowance of a ryegrass-dominated pasture

The effects of sward surface height (SSH) and daily herbage allowance (HA) on the defoliation pattern and grazing mechanics of early lactation dairy cows grazing on irrigated perennial ryegrass–white clover pasture were studied. The hypothesis tested was that SSH and HA affect intake and diet selection through their effects on the pattern of defoliation which is influenced by the resistance to prehension bites. Factorial combinations of two initial SSH (14 and 28 cm) and two daily HA (35 and 70 kg DM cow^?1 d^?1) were examined in a replicated experiment. The peak longitudinal tensile force required to break the sward portion encompassed in a 100 cm² area [bite fracture force (BFF₁₀₀)] was measured as an index of the resistance to prehension. The volume of herbage defoliated and herbage intake increased with SSH (P < 0·05) and HA (P < 0·01). Corresponding to an increase in HA from 35 to 70 kg DM cow^?1 d^?1, there was a proportional increase in the total defoliation area (TDA) and intake by 0·24 and 0·55 in the short sward compared with 0·16 and 0·32 in the tall sward respectively. The results of this experiment suggest that a consistent spatial pattern of reduction of the canopy exists during defoliation by cows and that the volume of sward canopy defoliated is the major variable affecting herbage intake. The BFF increased down the sward profile at a rate that was higher (P < 0·05) for the taller sward than for the shorter sward. It is proposed that a relatively lower resistance to prehension in the short sward compared with the tall sward explains the greater proportionate increase in TDA and intake corresponding to an increase in HA. The rate at which BFF₁₀₀ increases down the sward profile is suggested as a sward physical variable that can influence the defoliation process. The estimated time and energy costs of prehension bites are discussed in the context that larger bites are handled more efficiently than smaller bites.     

Spring and autumn animal treading effects on pre‐grazing herbage mass and tiller density on two contrasting pasture types in Ireland

A perennial ryegrass (Lolium perenne L.)‐dominated sward on a well‐drained soil (Experiment 1) and a creeping bent (Agrostis stolonifera L.)‐dominated sward on a poorly drained soil (Experiment 2) were subjected to four treading treatments: control (C, no damage), light damage (L), moderate damage (M) or severe damage (S) to quantify the effects on herbage dry‐matter (DM) production and tiller density. In Experiment 1, treading damage was imposed in spring. In Experiment 2, one‐third of the site was damaged in autumn, one‐third in spring and one‐third in both spring and autumn. Both sites were rotationally grazed after treading treatments. Pre‐grazing herbage mass was measured eight times in Experiment 1 and seven times in Experiment 2 on each plot, and tiller density was assessed four times in each experiment. In Experiment 1, pre‐grazing herbage mass was reduced by 30% in S plots at the first harvest after damage, but cumulative pre‐grazing herbage DM production was not different between treatments (12·7 t DM ha^?1). In Experiment 2, annual cumulative pre‐grazing herbage mass was reduced by between 14 and 49%, depending on intensity of treading damage event and season when damage occurred. Tiller density was not affected by treatment in either experiment. A perennial ryegrass‐dominated sward on a well‐drained soil was resilient to heavy treading damage. A creeping bent‐dominated sward on poorly drained soil requires a more careful grazing management approach to avoid major losses in cumulative pre‐grazing herbage mass production during wet weather grazing events.     

Pattern of herbage intake rate and bite dimensions of rotationally grazed dairy cows as sward height declines

To allow improved prediction of daily herbage intake of dairy cows in rotational grazing systems, intake behaviour was assessed throughout the day in 24‐h paddocks. Herbage intake in 16 lactating Holstein–Friesian cows was assessed using the short‐term (1‐h) weight gain method at four predetermined natural meal times throughout the day (early morning, T1; late morning, T2; mid‐afternoon, T3; and early evening, T4). The study comprised two 4‐day experiments, each with a cross‐over design of four blocks. In both experiments, cows grazed a 24‐h paddock daily, and the effect of the immediately previous grazing experience on intake behaviour was investigated throughout the day, taking account of daily fluctuations in the short‐term physiological condition of the cows. Experiment 1 was carried out to investigate overall grazing behaviour during meals as a sward is progressively depleted during the day, with intake being assessed within the paddock and, hence, on a depleted sward. Experiment 2 similarly investigated the effect of sward depletion and physiological condition throughout the day on intake, but cows were removed to fresh, undefoliated swards during intake measurement periods; thus, intake rate was not influenced by differences in sward condition. Intake behaviour from both experiments was compared to establish the effect on herbage intake of changes in sward state and non‐sward factors. In Experiment 1, sward surface height, available herbage mass, proportion of leaf and green leaf mass declined as the day progressed. Bite mass declined with sward depletion, and mean intake rate was 1·64 kg dry matter (DM) h^–1, which was significantly lower at T3 (P < 0·01) than during other meals. In Experiment 2, plot sward conditions did not change throughout the day, and intake behaviour also remained constant, with a mean intake rate of 2·11 kg DM h^–1. Mean bite depth as a proportion of pregrazing extended tiller height was constant throughout the day (mean 0·32). The results show that, although cows grazed throughout the day on progressively depleted swards, indicative of rotationally grazed paddocks (Experiment 1), bite mass declined linearly and intake behaviour was variable. However, where intake was assessed on high‐quality, undefoliated swards (Experiment 2), intake behaviour was similar regardless of the time of day and the immediately previous experience. There was some indication of an interaction between the effects of the sward and the physiological condition of the animal on herbage intake.     

Is there a tiller morphology ideotype for yield differences in perennial ryegrass (Lolium perenne L.)?

The objectives of this study were to measure the variability in ‘realized’ tiller morphology and identify whether there are emergent generalizations about a tiller ideotype for productivity or survival. Morphological traits during the vegetative growth stage were measured for 2 years in the field for eight perennial ryegrass cultivars: six diploid and two tetraploid cultivars under low and high nitrogen fertilizer (50 and 225 kg N ha^?1 year^?1 respectively). Traits measured were lamina width, length and area; pseudo‐stem length and diameter; an index of tiller shape; tiller dry weight; tiller density; and herbage mass. Almost all the traits differed significantly between cultivars and significant correlations were found between the 2 years. Principal component analysis identified that tiller morphology and dry‐matter yield were independent. No trait associations indicating alternative cultivar‐specific tiller morphologies were detected but cultivars differed in tiller size. The patterns of change between the 2 years indicated a possible morphological trajectory as swards age. The genetic compensatory relationship between tiller size and density had a 1:1 slope, indicative of constant yield. This implies that larger tillers would be more leafy, and higher leafiness of tetraploid over diploid cultivars was confirmed by the leaf:non‐leaf ratio and tiller shape index.     

Factors influencing tissue turnover during winter in perennial ryegrass-dominated swards

Despite low net grass growth rates in Ireland between mid-November and February, tissue continuously turns over as new leaves are produced and older leaves senesce. The effects of closing swards from grazing on three dates in autumn (1 September, 20 September and 10 October) on tissue turnover in a perennial ryegrass sward during the winter were examined at two sites in the north-east and south of Ireland. Leaf extension and senescence rates were greater in the south than in the north-east, and were greatest on the earliest closing date at both sites. Leaf appearance and extension rates at both sites were closely correlated with accumulated daily air temperature above 6°C and mean soil temperature. Site-specific factors also influenced tissue flux, possibly including grazing intensity at closing and density of tillers of perennial ryegrass. Swards in the autumn with high herbage masses [>2000 kg dry matter (DM) ha⁻¹ approximately] and high leaf area index (LAI; about 2·5 and above) had the greatest leaf senescence rates and the greatest declines in herbage mass during the winter. Leaf senescence rates per tiller in high LAI swards were correlated positively with amount of leaf per tiller and also the proportion of green leaf in the second youngest expanding leaf lost to senescence. As potentially a complex of interactive factors influences tissue flux in winter, it is proposed that the data from this study be used in conjunction with other data to construct a dynamic model to predict more reliably optimum closing date in autumn for herbage utilization in winter.     

Sodium fertilizer application to pasture. 8. Turnover and defoliation of leaf tissue

The effects of application of sodium fertilizer on the turnover and defoliation of leaf tissue were investigated in a perennial ryegrass (Lolium perenne) pasture grazed by dairy cows. Eight plots were allocated to treatments either with or without sodium fertilizer, with the former receiving 32 kg Na ha^–1 applied in five applications of NaNO₃ over the grazing season. An equivalent amount of nitrogen was given to the controls as ammonium nitrate, the application of which was reduced in the sodium treatment to equate nitrogen fertilizer applications for the two treatments. In nine periods between April and September, marked tillers were recorded to measure leaf turnover, leaf lamina growth and specific leaf weight and, when combined with tiller density measurements, gave an estimate of herbage flux for the sward. The defoliation and net growth of the marked tillers were monitored at 3-day intervals and the data were combined with tiller density and specific leaf weight data to determine the intake of the expanding, penultimate and oldest live leaf laminae. Sodium fertilizer application did not affect the rate at which leaves appeared, but it retarded their rate of disappearance. The extension rate and the specific weight of green laminae were both increased by sodium fertilizer application and therefore the net gravimetric growth rate was increased. Tiller density was not affected by sodium fertilizer application and hence the estimated herbage growth and net herbage flux were increased by sodium fertilizer application. Application of sodium fertilizer did not affect lamina length, and in both treatments the penultimate laminae were approximately twice as long as expanding and oldest live laminae. Defoliation frequency decreased from the expanding to the oldest live laminae in the control treatment without sodium. Sodium fertilizer application increased the frequency of defoliation of the oldest live leaf and also increased the length of the expanding leaf that was defoliated. For penultimate leaf laminae sodium fertilizer application reduced the defoliation frequency and length of foliage grazed. The dry-matter (DM) intake of the oldest live laminae was increased by the application of sodium fertilizer. It is concluded that sodium fertilizer application increases net herbage growth both by increased extension rate of leaf laminae and specific leaf weight and by delayed laminae senescence, and that it increases herbage DM intake by increasing the defoliation frequency of the oldest live leaf laminae.     

Effect of spring defoliation pattern on the mid‐season production and morphology of swards of perennial ryegrass cultivars of different maturity

The effect of three spring management treatments on the vertical distribution of dry‐matter (DM) yield and morphology of four cultivars of perennial ryegrass (Fennema, Corbet, Foxtrot, Melle) in mid‐season was investigated. The management treatments commenced with cuts on 15 February (Early), 1 March (Medium) and 29 March (Late), each followed by a 28‐day re‐growth period until the next cut and then further 21‐day re‐growth periods after each subsequent cut. This created four mid‐season measurement periods across the management treatments at cut 3 (5 April–17 May), cut 5 (17 May–28 June), cut 6 (7 June–19 July) and cut 7 (28 June–9 August). Tiller and sheath height and their ratio, and leaf lamina length, were measured prior to the four mid‐season cuts (cuts 3, 5, 6 and 7) when measurements of DM yield and proportions of leaf, stem and dead material in three herbage horizons (Lower, 0–8 cm; Middle, 8–15 cm; Upper, >15 cm) were made. There were significant responses in mid‐season to the management treatments involved complex interactions between management treatment and cutting date, which modified seasonal patterns in DM yield and leaf:stem ratio. There were significantly greater tiller heights, tiller:sheath ratios and leaf lamina lengths but lower sheath heights from the Early to Late management treatments. The greatest responses in morphological characteristics occurred in the Middle horizon compared with either the Lower (predominately stem and pseudo‐stem), or the Upper (predominately leafy) horizons. Distribution of DM yield between Middle and Lower horizons but not overall DM yields was significantly affected by management treatment. Morphological differences between cultivars were mostly in the Middle horizon and ranking of the cultivars was similar across the management treatments. The different responses of cultivars Fennema and Melle showed that genotype had a significant effect regardless of management. The leafiest mid‐season swards were achieved by delaying initial spring defoliation in the cultivar which had the lowest stem production.     

Incorporating grazing behaviour measurements in models to predict herbage intake by grazing dairy cows

Models to predict herbage intake were constructed using 168 dairy cow records from three grazing experiments. Variables included fell into three categories: animal state, sward state and animal behaviour. Linear regression models of varying complexity were obtained by removing variables from the best fitting model to reflect progressive lack of information availability on farms. Thus, behavioural variables were removed first, followed by sward surface height and milk fat concentration. Models were subject to outlier analysis and collinearity tests. Equivalent models were constructed using ridge regression to minimize collinearity problems. They were tested using 20 Holstein–Friesian dairy cows continuously stocked on a perennial ryegrass sward. A `best practice' treatment [7 cm sward surface height (SSH), 6 kg day⁻¹ concentrate (C)] was used together with treatments of SSH5/C6, SSH7/C8, SSH7/C0 and SSH9/C6. The best model accounted for 0.37 of the variance in the estimation data and contained the following variables: concentrate intake, milk yield, milk fat concentration, days in milk, sward surface height and chewing rate while ruminating. Model performance against test data was generally poor. This was mainly because of consistent underprediction of herbage intake, caused in part by the higher average herbage intakes in the test data compared with the estimation data.     

How does the vertical and horizontal structure of a perennial ryegrass and white clover sward influence grazing?

Mixtures of perennial ryegrass ( Lolium perenne L.) and white clover ( Trifolium repens L.) sown in alternate rows or in a thoroughly mixed matrix were grazed by sheep, either continuously or during short grazing tests, and were used to investigate the influence of the vertical and horizontal components of the sward structure on defoliation by sheep.
In an experiment under continuous grazing, the defoliation intensity was greater for white clover compared with perennial ryegrass leaves (0·80 and 0·58 respectively). In spring, perennial ryegrass leaves were more defoliated than white clover leaves, whereas the reverse was observed in summer. The ratio of the proportion of white clover to perennial ryegrass leaves grazed was negatively correlated with the difference between the surface height of the perennial ryegrass and white clover rows in spring. In both spring and summer, white clover leaves of the same extended leaf length had a higher proportion of them grazed than perennial ryegrass leaves.
In another experiment, during short grazing tests with perennial ryegrass–white clover swards that were grazed at the same sward surface height and at the same white clover content as in the previous experiment, there were no significant differences in the proportion of white clover and perennial ryegrass leaves grazed between strips of the two species and thoroughly mixed structures. The proportion of white clover leaves grazed was higher than that of perennial ryegrass leaves.
These results show that the differential defoliation by sheep of perennial ryegrass and white clover leaves varies according to their vertical distribution in the mixed canopy, but is little affected by their horizontal distribution. Even small differences in sward surface height between mixed perennial ryegrass and white clover can affect diet selection by sheep to a rather large extent.     

High perennial ryegrass seeding rates do not negatively impact pasture sward persistence

Poor persistence of perennial ryegrass swards is a common problem; however, there is a lack of long‐term studies to understand the mechanisms associated with poor persistence. This study describes an experiment to test the hypothesis that high ryegrass seeding rates (>18 kg seed per ha) reduce long‐term population persistence because of smaller plant size and poorer survival during the first year after sowing. Four cultivars, representing four functional types of perennial ryegrass, were sown at five seeding rates (equivalent to 6, 12, 18, 24 and 30 kg seed per ha) with white clover in three regions of New Zealand. Swards were monitored for 5 years. No evidence was found to indicate a lack of persistence of ryegrass‐based swards sown at higher seeding rates. During the first year, swards sown at higher seeding rates had greater herbage accumulation (except at the Waikato site), greater ryegrass tiller density and greater ryegrass content. This initial impact of high seeding rates had largely dissipated by the fourth year, resulting in swards with similar annual herbage accumulation, tiller density and botanical composition. Similarly, there were relatively few differences among cultivars for these variables. Although high seeding rates did not negatively impact sward persistence, geographical location did, with strong evidence of ryegrass population decline at the Waikato site for all treatment combinations, some decline in Northland, and stable populations in Canterbury. It is possible that productive perennial ryegrass pastures can only be sustained for 4–5 years in some situations, even when the best ryegrass technology and management practices are used.     

The relationship between level of sward height reduction in a rotationally grazed sward and short-term intake rates of dairy cows

The objective of this study was to investigate the relationship between level of sward height reduction (SHR) and short-term intake of herbage by lactating dairy cows offered swards differing in initial bulk density (BD). Three experiments were carried out in which cows were presented with swards representing different levels of SHR (nominally described as ungrazed, low, moderate and high). Experiments 1 and 2 differed with respect to initial sward BD [ungrazed sward 1·7 vs. 2·5 kg dry matter (DM) m^?3 respectively]. Experiment 3 investigated the interaction between BD and SHR. In each experiment, sixteen Holstein/Friesian cows (fitted with excreta collection bags) grazed for a 1-h period in 200-m² plots that had been grazed on the previous day to predetermined sward surface heights (SSH) (= levels of SHR). Herbage intake rates were assessed from changes in live weight, with a correction for insensible weight loss (IWL). Biting rates were recorded from visual observation. High levels of SHR were associated with a significant reduction in SSH, herbage mass and leaf fraction, and a significant increase in sward BD, stem and dead fraction, and DM concentration. Herbage intake, expressed either as DM intake per bite or DM intake per hour, declined as level of SHR increased from low to high. The level of SHR generally had no effect on biting rate. Intake rates varied from 1·9 to 4·4 kg DM h^?1, whereas DM intake bite^?1 ranged from 0·5 to 1·3 g. Pooled regression analysis identified SSH (P < 0·001; r² = 0·94) as the principal determinant of DM intake bite^?1. The regression equation was not significantly improved by the addition of terms for leaf fraction, BD, or herbage mass. In Experiment 3, a significant interaction between level of SHR and sward BD was observed. It is concluded that the principal factor controlling intake (g DM bite^?1 or kg DM h^?1), as swards are progressively grazed down, is SSH, but at a high level of SHR, sward BD also influences intake bite^?1.     

Effect of long-term changes in relative resource availability on dietary preference of grazing sheep for perennial ryegrass and white clover

Three replicate paddocks, each of 0·235 ha, containing adjacent monocultures of perennial ryegrass or white clover [50:50 by ground area, 6 cm sward surface height (SSH) at start of experiment] were continuously stocked with three yearling and four mature non-lactating, non-pregnant Scottish halfbred ewes for 12 weeks. Herbage intake, grazing behaviour and dietary selection were measured on seven occasions. Clover SSH declined rapidly over the first 5 weeks then stabilized at 1·2–1·6 cm, whereas perennial ryegrass SSH rose slightly initially, then declined gradually. Animals initially included proportionately c . 0·6 white clover in their diet but, by the end of the experiment, this had fallen to 0·3. Total daily herbage intake declined over the 12 weeks from 1·8 kg dry matter (DM) day^–1 at the start to 1·0 kg DM day^–1. Total grazing time increased from 561 min day^–1 to 649 min day^–1 at the end of the experiment. The results suggest that, despite overall herbage depletion and a greater depletion of white clover than perennial ryegrass as a result of the initial partial preference for white clover, the animals traded-off a reduced total intake and an increased grazing time in an attempt to maintain their initial preferred dietary composition.     

Response of herbage regrowth and water-soluble carbohydrate concentration of ryegrass species to defoliation practices when grown in a Mediterranean environment

Three experiments were conducted to determine the association between leaf number per tiller at defoliation, water‐soluble carbohydrate (WSC) concentration and herbage mass of juvenile ryegrass plants when grown in a Mediterranean environment. Seedlings of ryegrass were grown in nursery pots arranged side‐by‐side and located outside in the open‐air to simulate a mini‐sward in Experiments 1 and 2, and a mixture of annual ryegrass and subterranean clover (Trifolium subterraneum L.) was grown in a small plot field study in Experiment 3. Swards were defoliated mechanically with the onset of defoliation commencing within 28 d of germination. Frequency of defoliation ranged from one to nine leaves per tiller, whilst defoliation height ranged from 30 mm of pseudostem height that removed all leaf laminae in Experiment 1, to 50 mm of pseudostem height with some leaf laminae remaining post‐defoliation in Experiments 2 and 3. A positive relationship between herbage mass of ryegrass, WSC concentration and leaf number per tiller at defoliation was demonstrated in all experiments. In Experiment 1, the herbage mass of leaf, pseudostem and roots of tillers defoliated at one leaf per tiller was reduced to 0·10, 0·09 and 0·06 of those tillers defoliated less frequently at six leaves per tiller. However, the reduction in herbage mass from frequent defoliation was less severe in Experiment 2 and coincided with a 0·20 reduction in WSC concentration of pseudostem compared with 0·80 measured during Experiment 1. In Experiment 3, the highest harvested herbage mass of ryegrass occurred when defoliation was nine leaves per tiller. Although the harvested herbage from this sward contained senescent herbage, the in vitro dry‐matter digestibility of the harvested herbage did not differ significantly compared with the remaining treatments that had been defoliated more frequently. Leaf numbers of newly germinated ryegrass tillers in a Mediterranean environment were positively associated with WSC concentration of pseudostem and herbage mass. A minimum period of two to three leaf appearances was required to restore WSC concentrations to levels measured prior to defoliation thereby avoiding a significant reduction in herbage mass. However, maximum herbage mass of a mixed sward containing ryegrass and subterranean clover was achieved when defoliation was delayed to nine leaves per tiller.     

Seasonal changes in the crude protein concentration of mixed swards of white clover/perennial ryegrass grown without fertilizer N in an organic farming system in the United Kingdom

Changes in the crude protein (CP) concentration of white clover and perennial ryegrass herbage from a mixed sward were determined on six sampling dates from May to October in each of 2 years. The swards were grown without fertilizer N in an organic farming system and continuously grazed by dairy cows during the grazing season. The annual mean contents of white clover in the dry matter (DM) of the sward were 272·3 and 307·0 g kg⁻¹ in Years 1 and 2. The mean CP concentrations of the white clover and perennial ryegrass herbage were 251·6 and 151·9 g kg⁻¹ DM in Year 1 and 271·9 and 174·0 g kg⁻¹ DM in Year 2 respectively. The CP concentration of the white clover increased significantly during the grazing season from 220·0 to 284·1 g kg⁻¹ DM in Year 1 and from 269·0 to 315·5 g kg⁻¹ DM in Year 2. In the perennial ryegrass herbage the CP concentration increased from 112·2 to 172·6 g kg⁻¹ DM in Year 1 and from 142·7 to 239·5 g kg⁻¹ DM in Year 2. The rate of increase during the season in the CP concentration of the perennial ryegrass herbage was similar to the rate of increase recorded in the white clover herbage.