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.     

Effects of timing and intensity of defoliation on growth and development of white clover seedlings

The effects of different defoliation regimes on the growth and development of three contrasting white clover cultivars (S184, Menna and Alice) were assessed in three experiments in the glasshouse. Experiment 1, with clover growing on its own, investigated the effects of three times of onset × two intensities of defoliation. In Experiment 2, clover was grown with grass and there were two times of onset × two heights × two frequencies of defoliation. Experiment 3, also with grass, investigated the effects of changing defoliation frequency at different intervals from sowing. All clover cultivars responded similarly to the various treatments and there were no interactions between time of onset and subsequent defoliation regimes in Experiments 1 and 2. Without competition from grass (Experiment 1), defoliating early at the three leaf-stage of clover decreased the number of growing points by 32% and stolon weights and lengths by 50% compared with delaying defoliation until the nine leaf-stage. Maintaining one compared with two leaves per growing point had similar effects. Over 17 weeks undefoliated seedlings produced ten times more stolon than early defoliated or intensively defoliated seedlings. In competition with grass (Experiment 2) delaying defoliation significantly decreased all aspects of stolon growth. Seedlings growing in swards defoliated frequently and closely had most growing points whereas those defoliated infrequently had least. Stolon lengths and weights were larger for seedlings growing in swards defoliated frequently than for those defoliated infrequently at both heights of cutting. Mean weight of stolon per unit length was greater when swards were defoliated at 6 cm than at 2 cm height. Changing defoliation from every 2 weeks to every week (Experiment 3) decreased stolon growth slightly when the change was made early but increased it when the change was made late, although similar amounts of stolon were produced by seedlings continuously defoliated throughout every week and every 2 weeks. The results are discussed in relation to the seedlings' leaf complements and growth habit; the over-riding influence of grass competition is highlighted. The possibility of devising optimal defoliation strategies and the need to test these in the field are also outlined.     

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.     

The effects of defoliating grass in winter or spring on herbage yields and ensilage characteristics

Under Irish conditions, the digestibility in May of grass managed for silage production is sometimes lower than expected. In each of two successive years, replicate field plots were established to examine the effects of three defoliation heights (uncut or cut to a stubble height of 10 or 5 cm) applied in winter and/or spring on herbage yields harvested in May and again in July, and on chemical composition and conservation characteristics associated with first‐cut silage. Swards that were not defoliated in December or March had a dry‐matter (DM) yield and in vitro DM digestibility (DMD) in mid‐May of 6597 kg ha^?1 and 736 g kg^?1, respectively, in Year 1, and corresponding values of 7338 kg ha^?1 and 771 g kg^?1 in Year 2. Defoliating swards to 5 cm in December reduced (P < 0·001) May DM yields compared to swards that were not defoliated in both December and March, while herbage DMD in May increased (P < 0·001) when defoliated in December or March. There were no clear effects of defoliation height or its timing on herbage ensilability or resultant conservation efficiency characteristics. The effects of defoliation on July yield were the reverse of those observed for May, while the total yield of the December and March defoliations plus the two silage harvests increased as defoliation height was lowered in Year 2 only. It is concluded that defoliation in winter and/or spring can increase herbage digestibility but will likely reduce DM yields in May.     

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.     

Efficiencies of ryegrass and white clover herbage utilization in mixtures continuously grazed by sheep

A method for measuring the growth, senescence and defoliation fluxes in a mixture consisting of alternate rows of perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) was used to calculate the actual and potential efficiencies of grass and clover utilization under continuous sheep grazing. White clover contributed relatively more to the growth than to the herbage mass of the mixture and its relative growth rate was usually significantly greater than that of its companion grass. The primary reason for the greater potential efficiency of herbage use of the white clover component was, however, its lower rate of senescence. Greater potential efficiency was not reflected in a greater actual efficiency because the contribution of white clover to the total herbage removed was always smaller than its contribution to the growth flux of the mixture. Despite the commonly assumed sheep preference for white clover, the legume was usually the least defoliated species, presumably because of the lower surface height of white clover compared with grass, which resulted in a lower vertical availability of white clover, thereby restricting its defoliation rate. The positive net herbage accumulation observed with white clover at a constant sward surface height occurred partly through stolon and bud development and partly through the growth of undefoliated leaves from axillary growing points and contributed to a net herbage accumulation per unit dry weight in white clover greater than that in ryegrass. This resulted in an increase in the white clover content of the sward, which was observed independently by destructive measurements.     

Frequency of defoliation of native and naturalized species of the Flooding Pampas (Argentina)

This study describes the effect of herbage mass of species on defoliation patterns in a multispecies temperate–subhumid natural grassland of the Flooding Pampas at low, medium and high sward herbage mass [761  ±  8·7, 989  ±  137 and 1591  ±  44·0 kg green dry matter (DM) ha^–1 respectively] maintained by continuous grazing of cattle at variable stocking rates. Four native and naturalized warm-season species were studied: the grasses Cynodon dactylon , Leersia hexandra and Paspalum dilatatum and the legume Lotus tenuis . Sward state of the four species was described in terms of sward surface height, herbage biomass and the number of individual grass tillers and legume stems per unit area. The defoliation pattern was monitored twice-weekly during an experimental period of 3 weeks on permanently marked individual plants. Proportions of individual plants defoliated and the ranking of species selected differed among sward treatments and species. The results showed that the density of the species was a major factor determining the pattern of species defoliation and suggest that, in continuously grazed mixtures, the pattern of defoliation might be more dependent on the abundance of a species in the plant community than on species preferences. The results show the importance of considering this structural characteristic of swards in understanding the process of species selection by large herbivores in multispecies vegetation.     

The effect of height and frequency of mowing on the yield and composition of perennial ryegrass—white clover swards in the autumn to spring period

Ninety-six plots (3 × 2 m) of well-established perennial rye grass/white clover pasture were mown to heights of 2·7 (Low) or 3·96 (High) cm (rising plate meter) at 14-, 28-, 84- or 112-d intervals in autumn-winter. A 7-, 14- and 28-d mowing interval was superimposed in spring on each autumn–winter mowing interval treatment with the low and high mowing heights altered to 2·92 and 4·80 cm, respectively.
With the low cutting height, accumulated herbage DM was more than doubled (1806 ± 79 kg DM ha^-1) compared to a 'high' (754 ± 49 kg DM ha^-1) cutting height in autumn–winter and this was due to increased harvesting efficiency rather than growth as estimated by leaf extension. Although defoliation interval had no effect on DM yield, the grass component increased and clover decreased. The composition effect carried over into spring. On average, 3·5 tillers were produced over winter for each ryegrass tiller present in autumn and tiller densities were higher in spring. Tillers produced over autumn–winter contributed more than 60% of ryegrass growth by early spring.
In early spring (16–30 September), the low cutting height increased herbage DM yield, in mid-spring (1–14 October) it reduced DM yields particularly in combination with short defoliation intervals, while in late spring (14 October to 11 November) cutting height had no effect on DM yields.
Over the entire spring period there was a very marked effect of defoliation interval on DM yields.     

Effect of defoliation frequency and cutting height on growth,dry‐matter yield and nutritive value of Napier grass (Pennisetum purpureum (L.) Schumach)

Growth characteristics, dry‐matter (DM) yield, chemical components and in vitro dry‐matter digestibility (IVDMD) of Napier grass were studied in a randomized complete block design with three replications at Haramaya University, Ethiopia. The treatments were three defoliation frequencies (60, 90 and 120 d after Napier grass establishment) and five cutting heights (5, 10, 15, 20 and 25 cm above ground level). Except for leaf‐to‐stem ratio, all growth characteristics increased with decreasing frequency of defoliation. Similarly, DM yield of Napier grass increased as the frequency of defoliation decreased. Total ash, crude protein (CP), hemicellulose and IVDMD contents of Napier grass increased with increasing frequency of defoliation, whereas DM, acid detergent fibre, acid detergent lignin and cellulose contents increased with a decrease in defoliation frequency. In contrast, cutting heights had no significant effect on most chemical components and IVDMD contents, although total ash and CP decreased as the height of cutting increased. The CP yield per cut and per day was highest at an intermediate defoliation frequency. The results of this study indicated that Napier grass could be categorized under medium‐ to high‐quality herbage if defoliated at 90 d after establishment in the mid and highlands of Ethiopia.     

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.     

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.     

Influence of defoliation regime on herbage production and characteristics of intake by dairy cows as affected by grazing intensity

Three contrasting defoliation regimes for dairy cows—four cuts annually, severe rotational grazing and lax rotational grazing integrated with cutting—were compared in terms of herbage production, selection and intake per cow. Lax and severe grazing were compared simultaneously and grazing intensity was characterized by means of changes in herbage mass and sward height during grazing.
Herbage growth and yield were similar under cutting, severe grazing and lax grazing/cutting (120 t organic matter (OM) ha⁻¹). Herbage quality and leaf production were highest with severe grazing, which also had a less marked seasonal pattern of growth. The herbage intake per cow was 111 kg OM d⁻¹ with severe grazing and proportionately (0-20) higher at lax grazing/ cutting. 050 of the herbage yield was harvested by cutting at lax grazing/cutting. Selectivity was described with high certainly by the nutrient content prior to defoliation and by the degree of defoliation. Models to quantify this are presented.
Grazing intensity could be described by the size and degree of utilization of the fouled area, and herbage intake was quantified by means of herbage height before and after grazing. Within the range of 80-240 mm sward height prior to grazing, height measurements led to realistic and more accurate estimates than those obtained by measuring organic matter disappearance.
Herbage utilization was depressed significantly by increasing maturity of herbage due to lower nutritive value, but in particular due to reduced green leaf content and increased reproductive development. If sward height did not exceed 250 mm at any time, good utilization by grazing could be obtained.     

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.     

Effects of grass defoliation on the establishment and growth of slot-seeded white clover

White clover was slot-seeded into a low-fertility permanent pasture in May 1979. The effects of cutting interval (1, 2 and 4 weeks) and cutting height (3 and 7 cm), in the presence and absence of above-ground partitions, were recorded in terms of clover establishment and growth up to 15 weeks. The partitions resulted in a considerable increase in growth, stolon production and survival of clover plants, especially from 6 weeks after sowing, indicating a large effect of shoot competition from the surrounding sward. However, the effectiveness of defoliation in reducing grass competition appeared limited; close cutting did give some benefit to establishing clover but cutting frequency had little effect. In view of the limited success in reducing grass competition by cutting, further attempts at alleviating root competition are needed, possibly involving fertilizer placement and the use of grass-suppressing herbicides.     

Frequency and severity of defoliation of grass and clover by sheep at different stocking rates

The frequency and severity of defoliation of individual grass tillers and clover plant units was studied in Lolium perenne-Trifolium repens swards grazed by sheep at stocking rates ranging from 25 to 55 sheep ha^-1 and either receiving no N fertilizer or 200 kg N ha^-1. On average, sheep at the highest stocking rate defoliated individual tillers once every 4·2 d compared with once every 9·2 d at the lowest stocking rate with the removal of 58% and 47% of the leaf length of each tiller leaf at these stocking rates. Clover plant units were defoliated once every 4·2 d at the highest stocking rate and once every 7·2 d at the lowest stocking rate with the removal of 51% of its leaves and 12% of its stolon at the high stocking rate and 42% and 4% respectively at the low stocking rate. Differences in frequency and severity of defoliation between N fertilizer treatments were smaller than between stocking rates. Grass tillers and clover plant units were both defoliated less frequently and less severely in swards fertilized with N, though the difference in defoliation frequency between fertilizer treatments decreased as stocking rate increased. Defoliation frequency was related to the length of grass leaf per tiller or number of clover leaves per plant unit, and to the number of these tillers and the herbage on offer.     

Growth, colonization and productivity of two white clover cultivars strip-seeded into an upland Festuca-Agrostis sward

Small plots of a Festuca-Agrostis upland sward on a peaty gley podsol were strip-seeded during late June 1986 with white clover cvs Aberystwyth S184 or Menna at 4 kg ha⁻¹ and defoliated early (20 August) or late (3 September) and then frequently or infrequently (every 2 weeks or 4 weeks) until the end of September. All plots were defoliated in early November, at 3-weekly intervals during the growing season in 1987 and then grazed rotationally during 1988.
Satisfactory seedling establishment, representing 46% emergence, was achieved 5 weeks after sowing. The differential defoliation regimes had no persistent significant effects on clover development. S184 soon produced more leaves per seedling than Menna and a smaller proportion of its leaf number and weight were removed at each defoliation. Following large losses of leaves over the 1986–87 winter, SI84 had significantly more leaves per stolon than Menna; subsequently it also colonized the sward at a quicker rate. During 1987 amounts of herbage harvested (6.1 t ha⁻¹) were similar with the two clover cultivars, with S184 contributing 47% and Menna 44% of this respectively. SI84 made a larger contribution to yield during May and June but Menna was more productive during September and October. During 1988 clover populations were maintained with rotational grazing without additional fertilizer inputs.
The results show that, despite initial soil and climatic contraints, both small and medium-leaved clovers can be strip-seeded into upland swards with large subsequent benefits to yield and herbage quality. However, they also indicate the need for further experiments to determine the influence of sward morphology and defoliation regime on stolon branching rates and accumulation of growing points which, in turn, govern sward colonization.     

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 spring defoliation and fertilizer nitrogen on the growth of white clover in ryegrass/clover swards

An examination was made of the effects of different spring treatments on the growth of white clover in a ryegrass/white clover sward. Plots were either cut once (in February, March or April) or twice (in February and April) or left uncut. Nitrogen was applied to half of the plots in each instance. The clover was sampled at intervals of approximately 3 weeks from February to June to determine numbers of leaves and growing points and weights of plant parts. Rates of leaf appearance were also observed and estimates were made of total herbage mass from ground-level cuts.
Percentages of white clover in the herbage were higher in unfertilized than in fertilized plots and in defoliated than in undefoliated plots. The percentage increases that followed defoliation were usually maintained into later regrowth, showing that clover content was not automatically reduced as herbage mass increased. Increases in growing points were recorded after the beginning of April in defoliated unfertilized plots but not in undefoliated fertilized plots or in plots fertilized and defoliated twice during the spring period, in which numbers fell substantially.
Inverse relationships were found between rates of leaf appearance, or the number of green leaves retained per stolon, and herbage mass, whereas heights of clover and grass leaves and the percentage of dry matter allocated to petiole rather than leaf in the clover increased with increasing herbage mass.
We suggest that the observed differences between spring treatments in clover percentage result primarily from their differential effects on the formation and death of tillers and growing points in the early stages of regrowth.     

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.     

Effect of sowing date of triticale on seasonal herbage production in the central Appalachian Highlands of the United States

Triticale (X Triticosecale Wittmack) was evaluated as a complementary pasture to buffer those periods when herbage production from mixed perennial pasture is marginal in the central Appalachian Highlands of the United States. Triticale was sown every month from May to October for five consecutive years from 1999 to 2003. Plant population structure and herbage production were evaluated at intervals until May of the year following establishment. Triticale established quickly at all times of sowing except late October. Wet summers resulted in foliar disease and a rapid decline in plant density when triticale was sown in May and June. In contrast, during the relatively dry summer of 1999, triticale stands exhibited minimal decline. Triticale sown in August had a herbage yield of 1580 kg DM ha^?1 when harvested in October which was over twice the herbage yield of triticale sown in May, June and July. Average herbage yield in the following April of triticale sown in September was higher (1750 kg DM ha^?1) and less variable than herbage yields from other sowing dates. Plant and tiller populations declined throughout the following April but herbage yields in May were high due to stem and seed head development associated with reproductive growth. Incorporating areas of triticale into mixed‐species perennial pasture systems could buffer herbage production during hot and dry summer periods as well as during cool periods of late autumn and early spring.