Effect of defoliation management, based on leaf stage, on perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under dryland conditions. 1. Regrowth, tillering and water-soluble carbohydrate concentration

A field study was undertaken between April 2003 and May 2004 in southern Tasmania, Australia to quantify and compare changes in herbage productivity and water‐soluble carbohydrate (WSC) concentration of perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under a defoliation regime based on leaf regrowth stage. Defoliation interval was based on the time taken for two, three or four leaves per tiller to fully expand. Dry‐matter (DM) production and botanical composition were measured at every defoliation event; plant density, DM production per tiller, tiller numbers per plant and WSC concentration were measured bimonthly; and tiller initiation and death rates were monitored every 3 weeks. Species and defoliation interval had a significant effect (P < 0·05) on seasonal DM production. Prairie grass produced significantly more (P < 0·001) DM than cocksfoot and ryegrass (5·7 vs. 4·1 and 4·3 t DM ha^?1 respectively). Plants defoliated at the two‐leaf stage of regrowth produced significantly less DM than plants defoliated at the three‐ and four‐leaf stages, irrespective of species. Defoliation interval had no effect on plant persistence of any species during the first year of establishment, as measured by plant density and tiller number. However, more frequent defoliation was detrimental to the productivity of all species, most likely because of decreased WSC reserves. Results from this study confirmed that to maximize rates of regrowth, the recommended defoliation interval for prairie grass and cocksfoot is the four‐leaf stage, and for perennial ryegrass between the two and three‐leaf stages.     

Priority for allocation of water-soluble carbohydrate reserves during regrowth of Lolium perenne

A glasshouse study was undertaken to determine the priority within the perennial ryegrass (Lolium perenne L.) plant for leaf and root growth and daughter tiller initiation after defoliation, in relation to various levels of water-soluble carbohydrate (WSC) reserves at defoliation. Individual plants were arranged in mini-swards, and underwent varying defoliation frequencies and ambient temperatures before defoliation, and harvest heights at defoliation, to obtain a gradient of WSC content at H₁, the date when all plants were defoliated. Defoliation interval consisted of defoliating either three times at the one new leaf tiller^–1 stage (1-leaf stage) of regrowth, or once only at the 3-leaf stage, up to H₁, while night temperature in the week prior to H₁ was altered from 15°C to either 8 or 20°C. At H₁, plants were defoliated to a stubble height of either 20 or 50 mm. Plants were subsequently destructively harvested at days 4, 6, 8, 12, 18 and 27. Leaf and root extension and tiller dynamics were also measured. On a regrowth timescale, tiller initiation was most sensitive, root regrowth moderately sensitive, and leaf regrowth relatively insensitive to a decrease in WSC. The time of daughter tiller initiation also coincided with replenishment of stubble WSC levels. In contrast to this sequence of regrowth events following defoliation, the quantitative effects on growth were different, with elongation and survival of roots most affected by reduced WSC levels. A 30-fold difference in stubble WSC at H₁ between high and low WSC plants (1·52 vs. 0·05 mg tiller^–1) produced only a 4-fold increase in leaf dry matter (DM) after 27 d (2·2 vs. 0·6 g plant^?1), while tiller number plant^?1 increased 6-fold (138 vs. 23% increase in tiller number from H₁). Root elongation rate was 59 times higher in the high than in the low WSC plants (1·18 vs. 0·02 mm d^?1). From a pasture management perspective, the study confirms that defoliation, coinciding with the 3-leaf stage of regrowth and around a stubble height of 50 mm, optimizes persistence and productivity of perennial ryegrass. By allowing more rapid replenishment of WSC reserves, this optimal defoliation strategy enables a greater proportion of WSC to be allocated to maintain a more active root system, and promotes tillering, compared with more frequent and close defoliation.     

Effect of defoliation management, based on leaf stage, on perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under dryland conditions. 2. Nutritive value

A field experiment was undertaken between April 2003 and May 2004 in southern Tasmania, Australia, to quantify and compare changes in the nutritive value of perennial ryegrass (Lolium perenne L.), prairie grass (Bromus willdenowii Kunth.) and cocksfoot (Dactylis glomerata L.) under a defoliation regime based on stage of leaf regrowth. Defoliation interval was based on the time taken for two, three or four leaves per tiller to fully expand. At every defoliation event, samples were collected and analysed for acid‐detergent fibre (ADF), neutral‐detergent fibre (NDF) and total nitrogen (N) concentrations and to estimate metabolizable energy (ME) and digestible dry matter (DDM) concentrations. Amounts of crude protein (CP) and metabolizable energy (MJ) per hectare values were subsequently calculated. There was a significantly lower (P < 0·001) NDF concentration for perennial ryegrass compared with prairie grass and cocksfoot, and a significantly lower (P < 0·001) ADF concentration for cocksfoot compared with prairie grass and perennial ryegrass, regardless of defoliation interval. The CP concentration of cocksfoot was significantly greater (P < 0·001) compared with the CP concentrations of prairie grass and perennial ryegrass. The estimated ME concentrations in cocksfoot were high enough to satisfy the requirements of a lactating dairy cow, with defoliation at or before the four‐leaf stage maintaining ME concentrations between 10·7 and 10·9 MJ kg^?1 DM, and minimizing reproductive plant development. The ME concentrations of prairie grass (10·2–10·4 MJ kg^?1 DM) were significantly lower (P < 0·001) than for cocksfoot (as above) and perennial ryegrass (11·4–11·6 MJ kg^?1 DM) but a higher DM production per hectare resulted in prairie grass providing the greatest amounts of ME ha^?1.     

The importance of water-soluble carbohydrate reserves on regrowth and root growth of Lolium perenne (L.)

This glasshouse study aimed to determine the relative importance of water-soluble carbohydrates (WSC) and current photosynthate on root and top regrowth of perennial ryegrass (Lolium perenne L.). Individual plants were arranged in one of two miniswards (Experiments 1 and 2) and underwent varying defoliation frequencies designed to obtain a gradient of WSC content at the final harvest of each treatment (H₁), when all treatments were defoliated. In Experiment 1, the plants were defoliated either three times at the one new leaf per tiller stage of regrowth (treatment 3 × 1), once at the two-leaf and again at the one-leaf stage (treatment 2, 1), once at the one-leaf and again at the two-leaf stage (treatment 1, 2) or once only at the three-leaf stage (treatment 1 × 3), up to H₁. Leaf and root growth and other parameters were assessed over 6 d after H₁ in sunlight, and over a 4-week period in darkness, and related to initial plant WSC content. In Experiment 2, plant defoliation treatments were: 3 × 1, 1, 2 or 1 × 3. Leaf regrowth was assessed for 36 d until the plants had three fully expanded new leaves per tiller. Leaf regrowth in both experiments was significantly related to stubble WSC (below 50 mm height). In Experiment 1, plants were almost fully reliant upon plant reserves for the first 3 d of regrowth, with reliance decreasing up to 6 d. When regrowth of plants was compared after 1 week in light or in darkness, it was estimated that one-third of leaf regrowth was due to plant WSC reserves and the remainder due to photosynthesis. However, the capacity to photosynthesize and to grow roots after H₁ was also significantly related to stubble WSC content. In Experiment 2, there was a significant difference (P<0·01) between defoliation treatments on leaf dry matter (DM) yield at 12 d (×1 leaf tiller^?1) of regrowth, and this was, as in Experiment 1, significantly positively related to WSC content in the stubble. However, after 36 d of regrowth, DM yield of plants defoliated at 2 or 3 leaves tiller^?1 up to H₁ were similar, and both were significantly higher (P<0·01) than regrowth of plants defoliated at the one-leaf stage. After defoliation, the period of reliance on WSC reserves may be substantially increased in situations of shading (canopy competition or cloud cover) or if the new regrowth shoot is removed by regrazing.     

Changes in the physiology and feed quality of cocksfoot (Dactylis glomerata L.) during regrowth

Abstract A glasshouse study was undertaken to determine the physiological and morphological changes in cocksfoot (Dactylis glomerata L.) during regrowth after defoliation. Individual plants were arranged in a mini‐sward in a randomized complete block design. Treatments involved harvesting each time one new leaf had expanded (one‐leaf stage), up to the six‐leaf stage, with the plants separated into leaf, stubble (tiller bases) and roots. Stubble and root water‐soluble carbohydrate (WSC), stubble and leaf dry matter (DM), tiller number per plant and leaf quality (crude protein (CP), estimated metabolizable energy (ME) and mineral content) were measured to develop optimal defoliation management of cocksfoot‐based pastures. WSC concentration in stubble and roots was highest at the five‐ and six‐leaf stages. Mean WSC concentration (g kg^?1 DM) was greater in stubble than roots (32·7 ± 5·9 vs. 9·4 ± 1·5 respectively). There was a strong positive linear relationship between plant WSC concentration and leaf DM, root DM and tillers per plant after defoliation (Adj R² = 0·72, 0·88 and 0·95 respectively). Root DM plant^?1 and tiller DM tiller^?1 decreased immediately following defoliation and remained low until the three‐leaf stage, then increased from the four‐leaf stage. Tillers per plant remained stable until the four‐leaf stage, after which they increased (from 9·9 ± 0·5 to 15·7 ± 1·0 tillers plant^?1). Estimated metabolizable energy concentration (MJ kg^?1 DM) was significantly lower at the six‐leaf stage (11·01 ± 0·06) than at any previous leaf regrowth stage, whereas CP concentration (g kg^?1 DM) decreased with regrowth to the six‐leaf stage. Both the levels of ME and CP concentrations were indicative of a high quality forage throughout regrowth (11·37 ± 0·04 and 279 ± 8·0 for ME and CP respectively). Results from this study give a basis for determining appropriate criteria for grazing cocksfoot‐based pastures. The optimal defoliation interval for cocksfoot appears to be between the four‐ and five‐leaf stages of regrowth. Delaying defoliation to the four‐leaf stage allows time for replenishment of WSC reserves, resumption of root growth and an increase in tillering, and is before herbage is lost and quality falls due to onset of leaf senescence.     

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.     

Carbohydrate and crude protein fractions in perennial ryegrass as affected by defoliation frequency and nitrogen application rate

The objective of this study was to evaluate the effects of defoliation frequency (either at two‐ or three‐leaf stage) and nitrogen (N) application rate (0, 75, 150, 300, 450 kg N ha^?1 year^?1) on herbage carbohydrate and crude protein (CP) fractions, and the water‐soluble carbohydrate‐to‐protein ratio (WSC:CP) in perennial ryegrass swards. Crude protein fractions were analysed according to the Cornell carbohydrate and protein system. Carbohydrate fractions were analysed by ultra‐high‐performance liquid chromatography. Sward defoliation at two‐leaf stage increased the total CP, reduced the buffer‐soluble CP fractions and decreased carbohydrate fractions of herbage (P < 0·001). The effect of defoliation frequency was less marked during early spring and autumn (P < 0·001) than for the rest of the seasons. An increase in N application rate was negatively associated with WSC, fructans and neutral detergent fibre (P < 0·001), and positively associated with CP and nitrate (N‐NO₃) contents of herbage. Nitrogen application rate did not affect CP fractions of herbage (P > 0·05). The fluctuations in CP and WSC contents of herbage resulted in lower WSC:CP ratios during early spring and autumn (0·45:1 and 0·75:1 respectively) than in late spring (1·11:1). The herbage WSC:CP ratio was greater (P < 0·001) at the three‐leaf than the two‐leaf defoliation stage and declined as the N application increased in all seasons (P < 0·001). The results of this study indicate that CP and carbohydrate fractions of herbage can be manipulated by sward defoliation frequency and N application rate. The magnitude of these effects, however, may vary with the season.     

Interaction between water-soluble carbohydrate reserves and defoliation severity on the regrowth of perennial ryegrass (Lolium perenne L.)-dominant swards

A field-study was undertaken in Hamilton, New Zealand to determine if there was an interaction between water-soluble carbohydrate (WSC) reserve content and defoliation severity on the regrowth of perennial ryegrass-dominant swards during winter. Perennial ryegrass plants with either low or high WSC content were obtained by varying the defoliation frequency. At the third defoliation at the one-leaf stage and at the first defoliation at the three-leaf stage (harvest H₁), swards were mown with a rotary lawnmower to residual stubble heights of 20, 40 or 60 mm. All swards were then allowed to regrow to the three-leaf stage before again defoliating to their treatment residual stubble heights (H₂). Frequently defoliated plants contained proportionately between 0·37 and 0·48 less WSC in the stubble after defoliation, depending on the severity of defoliation. There was no interaction between WSC content and defoliation severity for herbage regrowth between harvests H₁ and H₂. Herbage regrowth was lower from swards containing low WSC plants compared with high WSC plants (2279 vs. 2007 kg DM ha⁻¹). Furthermore, swards defoliated to 20 or 40 mm had greater herbage regrowth compared with those defoliated to 60 mm (2266, 2249 and 1914 kg DM ha⁻¹ for swards defoliated to residual stubble heights of 20, 40 and 60 mm, respectively). Regrowth of perennial ryegrass was positively correlated with post-defoliation stubble WSC content within defoliation severity treatment, implying that WSC contributed to the defoliation frequency-derived difference in herbage yield. However, the effect of defoliation severity on herbage regrowth was not associated with post-defoliation stubble WSC content.     

Increased concentration of water-soluble carbohydrate in perennial ryegrass (Lolium perenne L.). Evaluation in dairy cows in early lactation

Twelve multiparous Holstein–Friesian dairy cows in early lactation were used to investigate the potential of using perennial ryegrass (Lolium perenne) with a high concentration of water‐soluble carbohydrates (WSC) to increase the efficiency of milk production. Ad libitum access to one of two varieties of zero‐grazed herbage was given continuously for 3 weeks: treatment High Sugar (HS), an experimental perennial ryegrass variety (Ba11353) bred to contain a high concentration of WSC, harvested in the afternoon; or Control, a standard variety of perennial ryegrass (cv. AberElan), harvested in the morning. All dairy cows also received 4 kg d^?1 of a standard dairy concentrate. Dairy cows given the HS diet treatment consumed 2·8 kg dry matter (DM) d^?1 more than Control dairy cows (P < 0·01), and the DM digestibility of the diet on the HS treatment was significantly greater than that of the diet on the Control treatment (0·75 vs. 0·72; s.e.d. 0·010; P < 0·05). Excretion of urinary purine derivatives (PD) tended (P < 0·1) to be higher from dairy cows on the HS treatment, implying increased microbial protein flow to the duodenum, although there was no significant difference in the apparent efficiency of rumen fermentation of either dietary nitrogen (N) or DM expressed as a ratio to urinary PD. Milk yields and milk composition were not significantly affected by dietary treatment, although true protein yields of milk were higher (P < 0·05) from dairy cows given the HS treatment. The proportion of dietary N excreted in urine was significantly lower from HS cows, although the values were low for both treatments (0·20 g g^?1 vs. 0·27 g g^?1; s.e.d. 0·020; P < 0·05). It is concluded that increased DM intakes by dairy cows given the HS treatment led to increased milk protein outputs. With a proportional decrease in urinary N excretion, the use of perennial ryegrass with a high WSC concentration, in the context of the harvesting regime used in this study, may help to reduce N pollution from dairy systems into which it is incorporated.     

Comparative defoliation tolerance of temperate perennial grasses

The defoliation tolerance of cultivars of four temperate perennial pasture grasses, perennial ryegrass (Lolium perenne, cv. Yatsyn1), phalaris (Phalaris aquatica cv. Australian), tall fescue (Festuca arundinaceae cv. Demeter) and cocksfoot (Dactylis glomerata cv. Porto), was determined under controlled conditions over a period of 12 weeks. Undefoliated plants were compared with defoliated plants, where only half of one leaf was left intact at the initial defoliation, and leaf regrowth was harvested every 3–4 d. The growth responses measured were plant tiller number, dry weight, relative leaf regrowth rate, root:shoot ratio, sheath:stem ratio and specific leaf weight. All species showed morphological adaptations that potentially increased their ability to tolerate defoliation (e.g. increased allocation to shoot at the expense of roots and lower specific leaf weights) but cocksfoot was found to be the most defoliation‐tolerant and perennial ryegrass the least. The adaptation that favoured cocksfoot most strongly was high sheath:stem ratio, which, it is proposed, allowed it to maintain photosynthesis and a level of carbon supply sufficient to support regrowth throughout the experiment. The strategy of perennial ryegrass which favours leaf growth and leads to rapid leaf turnover rates made it particularly susceptible to defoliation under the conditions of this experiment. This highlights the likely importance of defoliation‐avoidance responses in explaining the well‐known grazing resistance of this species. Phalaris and tall fescue showed responses that were intermediate between the other two species. The importance of defoliation‐avoidance mechanisms and implications for grazing management are discussed.     

Effects of continuous or rotational grazing of two perennial ryegrass varieties on the chemical composition of the herbage and the performance of finishing lambs

Plant breeding has developed perennial ryegrass varieties with increased concentrations of water‐soluble carbohydrates (WSCs) compared with conventional varieties. Water‐soluble carbohydrates are major metabolic and storage components in ryegrass. Therefore, if perennial ryegrass herbage is allowed to grow to greater heights it should contain higher water‐soluble carbohydrates concentrations, for example as under rotational grazing rather than continuous grazing by livestock. This study investigated this hypothesis and measured the performance of lambs grazed rotationally and continuously. Replicated plots of the variety AberDart (bred to express high WSC concentrations) or the variety Fennema were grazed by a core group of ten male Cheviot lambs for 10 weeks. Lambs were weighed and replicate forage samples were taken every 7 d. Concentrations of WSC in AberDart herbage were significantly (P < 0·05), but not substantially, higher than those in Fennema herbage. Rotational grazing did not increase the differential in WSC concentration between the AberDart and Fennema varieties. However, there was a tendency (P = 0·07) for lambs rotationally grazing the AberDart swards to have a higher final live weight than lambs grazing the Fennema swards. Overall, lamb performance was increased when either perennial ryegrass variety was rotationally rather than continuously grazed (P < 0·001).     

Leaf number as a criterion for determining defoliation time for Lolium perenne: 2. Effect of defoliation frequency and height

The object of this study was to determine the importance of frequency and height of defoliation on regrowth potential of Lolium perenne. Defoliation interval was based on stage of the regrowth cycle, as indicated by leaves per tiller.
Simulated swards of Lolium perenne cv Yatsyn were grown as individual plants in a glasshouse kept at a day/night temperature of 25°C/15°C.
Treatments imposed were defoliation at 2, 5 or 12 cm residual height, and low and high water soluble carbohydrate (WSC) level obtained by varying defoliation interval, i.e. defoliating at the 1-leaf or 3-leaf stage of the regrowth cycle. Regrowth after frequent short defoliations was only 65% of the less frequently defoliated plants taken over the full regrowth cycle. This was associated with a lower stubble WSC content (2·15 vs 17·5% in stubble) and a twenty-seven-fold difference in the amount of WSC in the stubble per plant. This difference in total WSC was a combined effect of more and heavier tillers and higher WSC content in stubble of plants defoliated less frequently at the end of the regrowth cycle. The regrowth of plants with WSC levels depleted by frequent defoliation when defoliated at 2 cm was significantly below that of those defoliated at 5 and 12 cm.
The results indicate the desirability of defoliating plants at the 3-leaf stage of the regrowth cycle. This not only allows the full regrowth potential to be expressed in that growth cycle, but also in the next cycle, by allowing the replenishment of WSC reserves and optimizing tiller status. The potential to regrow appears then to be based more on the total amount of WSC than the proportion of WSC in stubble.     

Increased concentration of water-soluble carbohydrate in perennial ryegrass (Lolium perenne L.): milk production from late-lactation dairy cows

Eight multiparous Holstein–Friesian dairy cows in late lactation were used to investigate the potential of using perennial ryegrass with a high concentration of water‐soluble carbohydrate (WSC) to increase the efficiency of milk production. After a pretreatment period on a common pasture, the cows were each given ad libitum access to one of two varieties of zero‐grazed grass continuously for 3 weeks. Treatments were: high sugar (HS), an experimental perennial ryegrass variety bred to contain high concentrations of WSC; or control, a standard variety of perennial ryegrass (cv. AberElan) containing typical concentrations of WSC. The two grass varieties were matched in terms of heading date. All animals also received 4 kg day^–1 standard dairy concentrate. Grass dry matter (DM) intake was not significantly different between treatments (11·6 vs. 10·7 kg DM day^–1; s.e.d. 0·95 for HS and control diets respectively), although DM digestibility was higher on the HS diet (0·71 vs. 0·64 g g^–1 DM; s.e.d. 0·23; P < 0·01) leading to higher digestible DM intakes for that diet. Milk yield from animals offered the HS diet was higher (15·3 vs. 12·6 kg day^–1; s.e.d. 0·87; P < 0·05) and, although milk constituent concentrations were unaffected by treatment, milk protein yields were significantly increased on the HS diet. The partitioning of feed N was significantly affected by diet, with more N from the HS diet being used for milk production (0·30 vs. 0·23 g milk N g^–1 feed N; s.e.d. 0·012; P < 0·01) and less being excreted in urine (0·25 vs. 0·35; s.e.d. 0·020; P < 0·01). In a separate experiment, using the same grasses harvested earlier in the season, the fractional rate of DM degradation, measured by in situ and gas production techniques, was higher for the HS grass than for the control. It is concluded that increased digestible DM intakes of the HS grass led to increased milk yields, whereas increased efficiency of utilization of the HS grass in the rumen resulted in the more efficient use of feed N for milk production and reduced N excretion.     

Effect of grazing management on herbage protein concentration,milk production and nitrogen excretion of dairy cows in mid‐lactation

The objective of this experiment was to use diurnal and temporal changes in herbage composition to create two pasture diets with contrasting ratios of water‐soluble carbohydrate (WSC) and crude protein (CP) and compare milk production and nitrogen‐use efficiency (NUE) of dairy cows. A grazing experiment using thirty‐six mid‐lactation Friesian x Jersey cows was conducted in late spring in Canterbury, New Zealand. Cows were offered mixed perennial ryegrass and white clover pastures either in the morning after a short 19‐day regrowth interval (SR AM) or in the afternoon after a long 35‐day regrowth interval (LR PM). Pasture treatments resulted in lower pasture mass and greater herbage CP concentration (187 vs. 171 g kg^?1 DM) in the SR AM compared with the LR PM but did not affect WSC (169 g kg^?1 DM) or the ratio of WSC/CP (1·0 g g^?1). Cows had similar apparent DM (17·5 kg DM cow^?1 d^?1) and N (501 g N cow^?1 d^?1) intake for both treatments. Compared with SR AM cows, LR PM cows had lower milk (18·5 vs. 21·2 kg cow^?1 d^?1), milk protein (0·69 vs. 0·81 kg cow^?1 d^?1) and milk solids (1·72 and 1·89 kg cow^?1 d^?1) yield. Urinary N concentration was increased in SR AM, but estimated N excretion and NUE for milk were similar for both treatments. Further studies are required to determine the effect of feeding times on diurnal variation in urine volume and N concentration under grazing to predict urination events with highest leaching risk.     

Competition in perennial ryegrass–white clover mixtures under cutting. 2. Leaf characteristics, light interception and dry-matter production during regrowth

The effect of defoliation interval on growth patterns of contrasting perennial ryegrass (Lolium perenne)–white clover (Trifolium repens) mixtures was studied. The dynamics of increase in leaf area, light interception and dry-matter (DM) production were measured within successive regrowth periods. No N fertilizer was applied. During 1995 six mixtures were cut eight (F1) or six times (F2) at a stubble height of 5 cm. The stubble composition was stable throughout the growing season: after harvest about 50 g DM m^?2 (with a white clover proportion of 0·52) was present with a leaf area index (LAI) of 0·5 (0·38 white clover). The percentage of intercepted radiation after cutting was 20–30% and increased during 3 weeks to about 95%. The relative growth rate of leaf area and DM was higher for white clover than for perennial ryegrass, with the proportion of clover in the LAI and DM increasing during each regrowth period. Mixtures with large-leaved white clover cv. Alice had a lower initial clover content after harvest, but a more rapid increase in clover LAI and DM than mixtures with the smaller leaved cvs Gwenda or Retor. Alice had the highest total and clover LAI and DM at harvest. Cutting frequency affected the change in white clover–perennial ryegrass ratio during regrowth. This was significantly higher in mixtures with Alice than in mixtures with Gwenda, but only under less frequent cutting (F2). In spring there was a mean white clover proportion of about 0·55 in the LAI and 0·45 in the total harvested DM. In summer the white clover proportion in the LAI and DM increased to 0·70–0·75. There was a decline during autumn, especially in F2 and in the mixtures with the small-leaved white clover cv. Gwenda and the medium-leaved cv. Retor. In contrast, grass DM and LAI declined from spring to summer. The decline in clover LAI in autumn was similar in Alice and Gwenda at frequent cutting (F1), but stronger in Gwenda in F2. Retor had the lowest clover specific leaf area (SLA). The SLA values of Alice and Gwenda were similar, SLA being similar between cutting treatments. No differences were found for leaf weight ratio (LWR) among the three white clover cultivars or between the grass cultivars, and LWR was not affected by cutting treatment. Defoliation interval had limited effects on the growth pattern and leaf characteristics of perennial ryegrass–white clover mixtures.     

Modelling the concentrations of nitrogen and water-soluble carbohydrates in grass herbage ingested by cattle under strip-grazing management

There is scope of increasing the nitrogen (N) efficiency of grazing cattle through manipulation of the energy and N concentrations in the herbage ingested. Because of asymmetric grazing by cattle between individual plant parts, it has not yet been established how this translates into the concentrations of N and water‐soluble carbohydrates (WSC) in the herbage ingested. A model is described with the objective of assessing the efficacy of individual tools in grassland management in manipulating the WSC and N concentrations of the herbage ingested by cattle under strip‐grazing management throughout the growing season. The model was calibrated and independently evaluated for early (April), mid‐ (June, regrowth phase) and late (September) parts of the growing season. There was a high correlation between predicted and observed WSC concentrations in the ingested herbage (R² = 0·78, P < 0·001). The correlation between predicted and observed neutral‐detergent fibre (NDF) concentrations in the ingested herbage was lower (R² = 0·49, P < 0·05) with a small absolute bias. Differences in the N concentration between laminae and sheaths, and between clean patches and fouled patches, were adequately simulated and it was concluded that the model could be used to assess the efficacy of grassland management tools for manipulating the WSC and N concentrations in the ingested herbage. Model application showed that reduced rates of application of N fertilizer and longer rotation lengths were effective tools for manipulating herbage quality in early and mid‐season. During the later part of the growing season, the large proportion of area affected by dung and urine reduced the effect of application rate of N fertilizer on herbage quality. In contrast, relative differences between high‐sugar and low‐sugar cultivars of perennial ryegrass were largest during this period. This suggests that high‐sugar cultivars may be an important tool in increasing N efficiency by cattle when risks of N losses to water bodies are largest. The model output showed that defoliation height affects the chemical composition of the ingested herbage of both the current and the subsequent grazing period.     

Patterns of leaf and root regrowth, and allocation of water-soluble carbohydrate reserves following defoliation of plants of prairie grass (Bromus willdenowii Kunth.)

This study utilized leaf stage‐based defoliation intervals to describe the concentrations and contents of water‐soluble carbohydrate (WSC) and nitrogen (N) in stubble and root reserves and their effect on the regrowth of prairie grass (Bromus willdenowii Kunth.) plants. The priority sequence for allocation of WSC reserves during the regrowth period was also investigated. There were substantially higher concentrations of WSC and N in the stubble compared with the roots following defoliation, confirming the stubble as the primary site for energy storage, with roots playing a lesser role. However, high R² values for the relationships between WSC concentration in roots and regrowth variables suggested that plants of prairie grass were reliant on WSC reserves from the roots in addition to the stubble to meet the energy requirements of plants until adequate photosynthetic tissue had been produced. The sequence of priority for allocation of WSC reserves followed the order of leaf growth, root growth and tillering during the regrowth period. Although WSC reserves were identified as the primary contributor to plant regrowth following defoliation, there was also a strong relationship between stubble N concentration and regrowth variables.     

Effects of continuous sheep stocking and strategic rest periods on the sward characteristics of binary perennial grass/white clover associations

Five binary perennial grass/white clover (Trifolium repens, cv. Menna) mixtures were evaluated over a 3-year period under continuous sheep stocking together with the imposition of a rest period for either an early or a late conservation cut; the experiment with plot sizes of 0·16 ha was replicated three times. The grass species and cultivars used were Merlinda tetraploid and Magella diploid perennial ryegrass (Lolium perenne), Prairial cocksfoot (Dactylis glomerata), Rossa meadow fescue (Festuca pratensis) and Goliath timothy (Phleum pratense). The greatest total lengths of white clover stolon developed in the meadow fescue (171·6 m m^?2) and timothy (151·9 m m^?2) associations compared with those in tetraploid perennial ryegrass (98·6 m m^?2), diploid perennial ryegrass (91·9 m m^?2) and cocksfoot (74·6 m m^?2) (s.e.d. 16·4, P < 0·001). On average, the proportion of white clover stolon that was buried was between 0·86 and 0·89 and this was more abundant in late than early season. Whereas timothy persisted, the persistence of meadow fescue was low under any of the managements tested and this was markedly reduced by the third grazing season. In the diploid perennial ryegrass sward, a late June to early August rest period for conservation enhanced white clover stolon length. An early April to late May rest period greatly reduced total white clover stolon length in both diploid perennial ryegrass and tetraploid perennial ryegrass associations (diploid perennial ryegrass-unrested 89 m m^?2, early rest 56·1 m m^?2, late rest 130·7 m m^?2; tetraploid perennial ryegrass - unrested 125·1 m m^?2, early rest 71 m m^?2, late rest 99·7 m m^?2; s.e.d. 19·19, P < 0·001). The numbers of white clover stolon growing points per unit stolon length were greatest when the sward was rested during late June to early August ?55·9 m^?1 stolon length compared with 45·7 m^?1 for an April to late May rest and 46 m^?1 in the absence of a rest (s.e.d. 2·59, P < 0·001). Likewise, the percentage of stolon above ground was greatest with the late June to early August rest ?15·78% compared with 10·61% for the April to late May rest and 7·69% for no rest (s.e.d. 1·569, P < 0·001). The complementary percentages of buried stolon indicate the important role this fraction has and the need to study stolon behaviour in grazing studies generally. It is concluded that, in relation to perennial ryegrass as a companion grass, meadow fescue and timothy allow better white clover development and cocksfoot less. However, other attributes have to be considered, for example the poor persistence of meadow fescue and the slower regrowth of timothy, both of which allow the invasion of weed grasses, or the lower acceptability of cocksfoot to livestock. The timing of the rest period before the conservation cut can influence white clover development considerably, but the effects differed with different companion grasses.     

The morphological and physiological responses of perennial ryegrass (Lolium perenne L.), cocksfoot (Dactylis glomerata L.) and tall fescue (Festuca arundinacea Schreb.; syn. Schedonorus phoenix Scop.) to variable water availability

There is a growing interest in the use of deficit irrigation and perennial pasture species other than perennial ryegrass (Lolium perenne L.) in temperate agriculture, in response to the decreasing availability of irrigation water. Deficit irrigation requires an understanding of plant responses to drought stress to ensure maximum dry‐matter return on water applied. A glasshouse study was undertaken to investigate some of the morphological and physiological responses of perennial ryegrass, cocksfoot (Dactylis glomerata L.) and tall fescue (Festuca arundinacea Schreb.; syn. Schedonorus phoenix Scop.) to varied moisture availability. One water treatment involved frequent applications of water to maintain a soil water potential of approximately ?10 kPa (100% treatment), and three other treatments involved applications at the same frequency, but using 33, 66 or 133% of the water applied in the 100% treatment. The water treatments continued over two plant regrowth cycles, followed by a ‘recovery’ phase of a single regrowth cycle during which all plants received the same water allocation as the 100% treatment. Depletion and replenishment of stubble water‐soluble carbohydrate (WSC) differed between the three species in response to soil moisture availability. By the second regrowth cycle, stubble WSC concentration and content in moisture‐stressed cocksfoot plants had increased, followed by a decrease during the subsequent recovery phase when the stored WSC reserves were utilized to support regrowth. The changes in stubble WSC reserves corresponded to the maintenance of relatively stable (i.e. the smallest reduction in leaf DM in response to moisture stress), but consistently lower DM production for cocksfoot compared with the other species. In contrast, moisture stress had no effect on the stubble WSC reserves of perennial ryegrass and tall fescue, with the exception of a significant decrease in WSC concentration under the 33% water treatment for perennial ryegrass. Perennial ryegrass achieved an intermediate DM yield and maintained positive growth rates throughout the study, even when watered at 33% of the requirement for optimal soil moisture levels. However, a more pronounced reduction in leaf DM in plants under moisture stress compared with the other species, combined with declining WSC reserves and the death of daughter tillers, highlighted the vulnerability of perennial ryegrass to poor persistence under prolonged drought conditions. Tall fescue appeared to have the greatest scope under moisture stress in terms of maintaining productivity and displaying attributes that contribute to persistence. Its leaf DM was consistently greater than that of the other species, displaying a smaller decline in growth under water stress compared to perennial ryegrass and an ability to recover faster upon re‐watering. This study has expanded the information available that compares and defines the potential of each species under moisture stress and emphasizes the importance of balancing short‐term DM production with long‐term persistence in choice of pasture species.     

Effect of plant age and severity of defoliation on regrowth of sheep's burnet during substrate moisture depletion

Regrowth of 3- and 4-month old (‘young’ and ‘old’ respectively) sheep's burnet [Sanguisorba minor ssp. muricata (Spach) Briq.] was studied under limiting and non-limiting moisture conditions in a glasshouse. Moisture deficits were imposed by using a single cycle of withholding moisture until first wilting. Plants of each age were defoliated severely at three levels which represented approximately 80–100% canopy removal. These levels were based on the proportion retained of the eight most mature leaves on each plant and were referred to as complete [0% residual leaf area (rLA)] and partial [50% rLA (four leaves) and 100% rLA (eight leaves)] defoliation. Vegetative growth and total non-structural carbohydrate (TNC) levels were studied. Leaf number (0-8 leaves), area (0-115 cm²) and dry weight (0-1·0 g) differed (P < 0·05) between defoliation intensities at the start of regrowth, while stubble (1·2 g) and root (12.·6 g) dry weights were similar. Soluble sugars [< 6% dry matter (DM)] and starch (< 1% DM) occurred in leaf, stubble and root. Old plants were morphologically and physiologically more developed than young plants. For example, stubble (2·0 g) and root (21·5 g) dry weights of old plants were greater (P < 0·05) than those of young plants (04 and 3·7 g respectively). Defoliation intensity had a major effect on regrowth, with completely defoliated plants at the final harvest having leaf numbers (forty-nine leaves) and areas (235 cm²) almost twice those of partially defoliated plants. Stubble soluble sugar levels (38% DM) were lower than those of partially defoliated plants (5·5% DM), and it was suggested that these contributed to regrowth. Moisture regime had a negligible influence on plant growth. However, plants in the dry regime had soluble sugar levels 1·4 (stubble) -1·7 (roots) times higher than those watered adequately, which suggested that plants adjusted to the water depletion. The effects of plant age on regrowth were similar for most characters, but the larger and physiologically more mature old plants would probably be more tolerant of successive defoliations.