Growth and photosynthesis of tall and short cultivars of white clover with tall and short grasses

A comparison was made of canopy structure, photosynthesis and dry weight production, during growth periods in spring and summer, of a large-leaved and a small-leaved cultivar of white clover (Blanca and S184), grown in mixtures with perennial ryegrass or tall fescue. Both clovers had leaves in the topmost, best-lit layer of the canopies, even when sward surface height reached 40 cm in early June. Except early in the spring growth period, both clovers had a greater proportion of their leaf area near the top of the canopy than did their companion grass. Both clovers had a greater relative growth rate than the grass, increasing their percentage of the total crop dry weight during both spring and summer growth periods. Differences between clover cultivars were relatively minor and did not affect their dry weight in either measurement period. During the first growth period there was more clover dry weight with tall fescue than with ryegrass. There were no effects of clover cultivar on grass dry weights.     

SOME SELECTIVE EFFECTS OPERATING ON WHITE- AND RED-CLOVER IN SWARDS

Populations of white- and red-clover were examined after they had been grown for several years in grass/clover swards. The method of sowing, nitrogen application, management and season affected the ratio of very similar clover cultivars differing specifically in the leaf mark when sown in the same mixture. The companion grass also infiuenced the proportion of legume cultivars in the sward. In addition, the mean date of emergence of a single cultivar of broad-red clover was affected by management.     

CLOVER NITROGEN AND SWARD PRODUCTIVITY

Under mowing conditions in the absence of the animal, ultra-simple swards based either on ryegrass or cocksfoot gave large increases in yield of dry matter and nitrogen from the inclusion of white clover in the seeds mixture. Raising the clover seed rate from 1 to 3 lb./acre gave a further response in yield.
As the swards aged, the continuing effect of increased clover seed rate was dependent on applied nitrogen.
The rate of transference of nitrogen from clover to the soil depended upon the species of the associated grass, the clover seed rate, the age of the sward and the degree of reduction of the clover in the sward.
Suppression of clover led to an increase in soil nitrogen, particularly with high seed rates of clover.
Dry matter response to applied nitrogen (69 lb./acre/annum) was greater in cocksfoot than in ryegrass. Fertilizer nitrogen had little effect on the yield of nitrogen from the sward except where the high seed rate of clover had been used.     

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.     

Diet selection by sheep and goats and sward composition changes in a ryegrass/white clover sward previously grazed by cattle, sheep or goats

An experiment was conducted to examine how variation in the composition and structure of mixed grass/white clover swards affected diet selection by sheep and goats. Sward composition in a mixed perennial ryegrass/white clover sward was manipulated by continuous grazing from 28 May to 28 July (pre-experimental phase) with cattle, sheep or goats, and then from 29 July to 2 September (experimental phase) with sheep or goats in a factorial design replicated twice. Sward surface height was maintained at 6 cm by regular adjustment of stocking density. Grazing by different sequences of animal species resulted in significant differences in the proportions of white clover in the sward, and especially in the proportion of clover lamina and petiole. Grazing by goats in the pre-experimental phase led to greater proportions of clover lamina and petiole in the whole sward and the sward surface. The proportion of white clover in the diet selected by sheep in the experimental phase was consistently higher than that in the sward as a whole, but was closely related to that near to the sward surface (approximately the top 2 cm). For goats there was no significant relationship between the proportion of clover in the diet and in the whole sward, and they generally selected a diet with a lower proportion of white clover than was present in approximately the top 2 cm of the sward. It is concluded that on mixed grass/white clover swards goats do not graze as deeply into the sward as sheep and that this results in a lower proportion of white clover in their diet and therefore allows higher proportions of white clover to develop under grazing by goats than by sheep.     

Growth of grass/clover mixtures during winter

The percentage of clover in a mixture of Huia white clover and Melle perennial ryegrass decreased during each of three winters. In the most severe winter, both grass and clover suffered a net loss of weight, clover losing two–thirds of its maximum weight while grass lost about one–third. In the mildest winter, grass continued to gain weight throughout, while clover lost weight. Such losses, when combined with conditions such as the use of nitrogen fertilizer, which prevent a recovery of clover percentage during summer, will lead to a long–term decrease in the clover content of a sward.
The losses from the clover were contributed to by all plant parts especially leaf, which lost between 60% and 95% of lamina weight. The change from autumn to winter caused a greater decrease in the number of live leaves per shoot. and in the area and weight of individual leaves, in clover than in grass. Other work has shown that there is a greater decrease in photosynthesis in clover than in grass, probably owing to changes in canopy structure; this is one cause of clover's poor performance, though differences between the two species in the effect of winter on the distribution of the carbon fixed in photosynthesis and in losses of weight in respiration and tissue death may also be involved.     

Effects of interval between harvests and application of fertilizer N in spring on the growth of perennial ryegrass in a grass/white clover sward

A perennial ryegrass/white clover sward, which had been grazed for over 2 years, was cut at 1-, 2-, 3- or 6-week intervals from 18 April to 28 November 1986. Two rates of fertilizer N application in spring, 0 and 66 kg N ha⁻¹, were compared. Perennial ryegrass growth was studied in three 6-week periods, beginning on 18 April, 18 July and 17 October. Clover growth was studied in the same three periods and described by Fisher and Wilman (1995) Grass and Forage Science , 50 , 162–171.
Applied N increased the number of ryegrass tillers m⁻², the rate of leaf extension and the weight of new leaf produced tiller⁻¹ and m⁻²per week. Increasing the interval between cuts reduced the number of ryegrass tillers m⁻² and increased the rate of leaf extension, weight tiller⁻¹and the weight of new leaf produced tiller⁻¹week⁻¹. Increasing the interval between cuts increased the weight of new ryegrass leaf produced m⁻² where N had recently been applied, but otherwise had little effect on the weight of new leaf produced m⁻². Applying N favoured the grass in competition with the clover in every respect, whereas increasing the interval between cuts only favoured the grass, compared with clover, where N had recently been applied; where N had not been applied, the ratio of ryegrass tillers to clover growing points in the sward was very little affected by the interval between cuts.     

A comparison of grass/white clover and grass silages offered to dairy cows as the sole feed

Two diets were compared: perennial ryegrass ( Lolium perenne L.) silage and perennial ryegrass/white clover ( Trifolium repens L.) silage, in each case fed ad libitum , without supplementation, to lactating dairy cows. The comparison was made with silages cut on each of four dates. The crops were wilted to a dry matter content of at least 256g kg^-1 and no additives were used.
The grass/clover silages were well preserved at all four cuts. The grass was well preserved at three cuts, but, at a May cut, the grass silage was less well preserved and less digestible than the grass/clover silage. When the May silages were fed, milk yield was higher with grass/clover than with grass. Taking the experiment as a whole, however, milk yield and composition were similar on the two diets. The grass/clover silages had a lower proportion of cell wall and their intake by dairy cows was consistently higher than that of the grass silages.
It is concluded that perennial ryegrass/white clover crops can be ensiled successfully and fed successfully, with high intake, to lactating dairy cows, but it should not be assumed that cows will give more milk than when fed equivalent all-grass silage.     

The diet ingested by sheep grazing swards differing in white clover and perennial ryegrass content

A series of twenty-four swards containing different proportions of white clover (0·20-0·25) and perennial ryegrass were created by using different seed mixtures, herbicide applications and previous cutting Frequencies. These swards were used to study the diet of oesophageally-fistulated wether sheep which grazed the various swards for a 30-min period after 1, 2 and 3 weeks of regrowth.
The proportion of white clover in the diet was generally greater than that in the sward. Fifty-seven percent of the variation in the proportion of white clover in the diet could be attributed to the proportion of white clover in the sward. White clover and perennial ryegrass leaf and stem were grazed to the same height and the proportion of white clover in the grazed horizon of the sward explained 83% of the variation in the proportion of white clover in the diet. The proportion of white clover in the diet was greater than the proportion in the grazed horizon of the sward in week 3 of regrowth, but not in weeks 1 and 2, and greater when the proportion of white clover in the grazed horizon was lower than 0·20. Both these observations were interpreted as indicating selection for white clover by the sheep within the grazed horizon.
There was a positive and linear relationship between the depth of the grazed horizon and sward height which, together with the relationship between the proportion of white clover in the grazed horizon and in the diet, would allow the prediction of the proportion of white clover of the diet from the height and the white clover content of the grazed horizon of the sward.     

The effects of variety, cutting interval and nitrogen application on the morphology and development of stolons and leaves of white clover

Six varieties of white clover, each grown with perennial ryegrass, four intervals between cuts and two levels of applied nitrogen in all combinations, were compared in a field experiment during the first 27 months after sowing. Information about yields, crop fractions, heights and ryegrass tillers has been presented in an earlier paper (Wilman and Asiegbu, 1982). The present paper is concerned with the more detailed studies of white clover, which help to explain the yield results and contribute to the understanding of the response of this species to management when grown in competition with grass. Increasing the interval between harvests increased the length of clover stolon per unit area of ground and increased stolon diameter, petiole length, weight per leaf and number of leaves harvested as a proportion of the number present in the sward while only slightly reducing the rate of leaf emergence, helping to explain the positive effect of increasing the interval on clover yield noted in the earlier paper. During regrowth, successive leaves had longer petioles and the length of individual petioles increased beyond the stage at which the leaflets were fully opened. Weight per leaf in clover increased considerably from April to June and declined to below the April value by October. It was shown that weight per leaf can be greatly increased by increasing the interval between harvests without reducing the number of leaves harvested per unit area per year. The stolon length measurements provided some support for the view that medium large-leaved varieties of white clover can with advantage be defoliated rather less frequently than small-leaved varieties. Stolon length was less adversely affected by applied N in the medium large- than in the small-leaved varieties. The small-leaved varieties had thinner stolons than the medium large-leaved varieties but about twice the stolon length when no N was applied, and a relatively high proportion of leaves which escaped defoliation. The application of N reduced stolon diameter, increased petiole length and had little or no effect on weight per clover leaf.     

Short-term effects of nitrogen on the growth and nitrogen nutrition of small swards of white clover and perennial ryegrass in spring

Small swards of white clover and perennial ryegrass were established in Perlite in a heated glasshouse, as either monocultures or mixtures of equal plant numbers. On 26th March, 1984 the swards were moved outside and their growth studied over the period to 29th May. All swards received a basal level of nitrate N to simulate soil mineralization and two-thirds received additionally the equivalent of 80 kg N ha⁻¹. Over the experimental period (26th March-29th May) clover maintained its proportion of total mixture dry weight In swards given 'fertilizer'-N and increased its proportion in those given only 'basal' N, However, clover declined as a percentage of total mixture dry weight during the first period of the experiment when total leaf area index (LAI) and ambient temperatures were low, and increased its percentage later when temperature and LAI had risen. Changes in tiller and stolon growing point numbers were not good indicators of changes in dry weight. Relative yields and relative replacement rates also were not good indicators of relative performance in mixture. Where 'fertilizer'-N was applied, clover derived less of its nitrogen from that source than grass although their uptakes per unit shoot dry weight were similar.     

The effect of sward height on responses of mini-swards of perennial ryegrass/white clover to slurry application

The influence of sward height at the time of slurry application on sward responses to slurry was investigated using perennial ryegrass ( Lolium perenne )/white clover ( Trifolium repens ) mini-swards under greenhouse conditions. Pig slurry, cattle slurry and a fertilizer control were applied either to: swards cut 1 d before slurry application to heights of 2, 4 or 8 cm (CH2, CH4 or CH8); or to swards cut to 4 cm with regrowth intervals of 1, 4 or 8 d before slurry application (RI1, RI4 and RI8). Scorch, smother and growth of marked clover stolons and grass tillers were monitored after slurry application. Dry-matter yields of both species were recorded over two harvests. Electrolyte leakage from leaves was used to assess leaf damage. Both slurries increased leakage, which was greatest from clover leaves, with biggest increases caused by pig slurry. Yield responses to slurry application varied with plant species and sward height. Reduced grass growth after slurry application to long swards (CH8, RI4, RI8) was probably caused by greater smother. In short swards (CH2), grass leaf growth was not depressed following slurry application but grass yields were reduced. Ammonia volatilization losses may have been greater in both long and short swards, contributing to yield reductions. Yield responses to slurry were greatest in swards cut to 4 cm, 1 d before application.     

RESPONSE OF GRASSLAND TO NITROGENOUS FERTILIZER IN THE WEST OF ENGLAND

Extensive replicated plot experiments were carried out at Henley Manor Farm from 1956–1961 to obtain accurate measurements of herbage production and response to nitrogenous fertilizers under West of England conditions. 356 plots were used; use of the herbage by both cutting and grazing was studied. The principal results are: Grass/ clover swards (containing approx. 35% clover) grown without nitrogenous fertilizer averaged 75 cwt dry matter per acre per annum. Application of nitro-chalk to initially grass/clover swards reduced the clover content of the herbage. About 133 lb N/annum was required by an all-grass sward to equal the production of a grass/clover sward without N. Attempts to increase production of grass/clover swards by using N for early growth and relying on clover for mid-season growth were unsuccessful in 2 out of 3 years. Swards (initially grass/clover) gave significant increases in total dry-matter production from regular use of the lowest level of N (26.0 or 34.7 lb N/acre/cut or graze) in 44 out of 49 cases. Extremely high yields were obtained from the heaviest use of N (104.2 lb N/acre/silage cut). 52.1 lb N/acre/cut or graze was the optimum rate of application on ail-grass and grass/clover swards. Up to about 350 lb N/acre/annum the dosage-response curve was very nearly straight for all-grass and grass/clover swards. On the grazed plots herbage left ungrazed amounted on average to only 6% of the total. The health of all stock on the high nitrogen plots (as well as all others) was excellent.     

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.     

Slot-seeding investigations 7. The effects of two grasssuppressants on the growth of white clover introduced into permanent pasture

The benefits of reducing competition from the surrounding grass sward against slot-seeded clover were measured in terms of white clover dry matter production following applications of two grass-suppressing herbicides, propyzamide and carbetamide.
Clover was slot-seeded in early June 1983 and herbicides were applied in September and October 1983. The later application was more damaging to the surrounding sward. Generally, the September spraying was more effective in encouraging white clover, especially from July of the following year. Only by September 1984 did the October applications result in higher clover yields than the September applications.
Propyzamide had the greater stimulatory effect on white clover production but it also damaged the grass more than carbetamide. Increasing the dose of each herbicide increased grass damage. However, with treatments applied in September, the loss was more than compensated for by an increase in clover and in total dry matter production at the end of the year.
The results suggest that either of these herbicides, but especially propyzamide, can be used to increase both total dry matter production from the sward and the contribution from slot-seeded white clover.     

Effect of interval between harvests and spring-applied fertilizer N on the growth of white clover in a mixed sward

A perennial ryegrass ( Lolium perenne )-white clover ( Trifollum repens ) sward, that had been grazed for over 2 years, was cut at 1-, 2-, 3- or 6-week intervals from 18 April to 28 November 1986. The effects of two rates of N application in spring, 0 and 66kg Nha⁻¹, were compared. Clover growth was studied in three six-week periods that began on 18 April, 18 July and 17 October respectively. Increasing the interval between cuts increased the yield of herbage without reducing the proportion of clover in the harvested herbage, but the combination of applied N and the six-week interval was harmful to clover production. Increasing the interval between cuts tended to increase the proportion of resources allocated to stolons rather than to leaves, and to increase the weight of new dry matter (DM) per growing point rather than per m². In the third period of the study, when the ten youngest internodes per stolon were examined separately, all ten ages of internode were found to have been affected by the cutting management. This was indicated by the positive effect of the length of the interval between cuts on the weights of N, P and K per intemode. The concentrations of N, P and K were highest in the youngest intemodes. The application of N reduced the proportion of new DM allocated to stolen rather than to leaf and it reduced the number of clover growing points per m².     

Influence of initial cutting date and cutting frequency on yield and quality of star, elephant and Guinea grasses

Two experiments were conducted to estimate the influence of initial cutting dates in March and April and of cutting frequencies on Guinea grass, elephant grass and star grass productivity. Increasing delay in date of first cut in the season resulted in a progressive increase in dry-matter yield. 72–81% of the tiller population at any cut were vegetative and this helped to maintain good aftermath yield for each date of first cut. Early April cuts gave the highest dry-matter yield and early March the least.
Harvesting frequencies affected the sward productivity such that the annual dry matter yield increased with increasing harvesting interval from 6800 kg ha^-1 for a 3-week interval to 13,000 kg ha^-1 for a 10-week interval. The proportion of green leaf in the dry matter dropped from 57·7% at 3-week intervals to 32·0% at 10-week intervals. Seasonal effects showed that potential yield was reduced by a short interval between harvests early in the season but not late in the season. Species differences in quality and in the harvesting interval that gave the maximum yields were noted. These results are discussed in relation to management of the sward throughout the growing season.     

Contribution of white clover varieties to total sward production under typical farm management

White clover varieties, potentially suitable for inclusion in seed mixtures for mixed stock-rearing farm systems, were evaluated when growing with S23 perennial ryegrass under rotational sheep grazing with a silage cut in late May or early June, as practiced on farms. Monoculture grass swards were also included to enable the direct and indirect contribution of white clover to total sward production to be evaluated.
White clover increased total sward production during all three years of the trial by an average of 50%. Although clover content of swards were similar, large differences occurred in their grass content, especially in the third year, when difference in total yields of swards based on large–leaved clover varieties was 2 t ha ⁻¹ while difference in clover yield was only 0-6 to ha^–1 The indirect contribution of white clover, namely the extra grass resulting from N transfer, was greater in the spring than in the autumn. It was also greater for Nesta than for other varieties. and exceptional for this variety in that the increase in grass yield above that of grass monoculture was maintained over three harvest years.     

Seasonal variation in white clover content and nitrogen fixing (acetylene reducing) activity in a cut upland sward

The seasonal variation in herbage mass and nitrogen fixing (acetylene reducing) activity of white clover in an upland sward, cut weekly to 3·5 cm from mid-May until mid-October, was measured. Acetylene reducing activity (ARA) was measured over a 24-h period at 3-weekiy intervals starting on 3 March 1983. Clover leaf and Stalon biomass was measured by harvest of the assay truces, and from mid-May quadrat euts to 3·5 cm above the soil surface provided estimates of herbage accumulation.
Little A RA was detected in March, but activity increased substantially after 10 cm soil temperatures reached > 3°C, and peak activity per unit of clover leaf dry weight occurred in June and July; standing clover leaf dry matter increased during the season to a maximum of 60·5 g m⁻¹ in June. Acetylene reducing activity was positively correlated with the number of rooted nodes, and Stalon and leaf dry weights in early spring. Thereafter, except during a period of summer drought, ARA was positively correlated with the amount of clover leaf material.
Clover population density increased during the season and maximum growing point numbers (5540 m⁻¹) occurred in September; maximum leaf number per unit area (12 984 m⁻¹) was found in October, prior to the final cutting of the site.
Results suggest that higher levels of nitrogen fixation in upland swards should be obtained if sward management regimes, which encourage a high clover leaf area, are adopted.     

The effects of contrasting cutting regimes on the components of clover and grass growth in microswards

Microswards of white clover and perennial ryegrass were subjected to one of four treatments: weekly cutting to 3·5 cm, weekly cutting incorporating a period of no cutting for 6 weeks starting 27 April (early rest), weekly cutting incorporating a period of no cutting for 6 weeks starting 8 June (late rest), or cutting every 3 weeks. Two sward types were used: a mixture of white clover cv. Milkanova with perennial ryegrass cv. Melle, and white clover cv. Kent with perennial ryegrass cv. Melle. Growth measurements (leaf appearance, branching/tillering and stolon internode length) were confined to the first three treatments with records collected during contiguous 21-d measurement periods. Vertical height increments of clover and grass and red:far-red light ratios at the sward bases were also recorded at frequent intervals. At the end of the experiment population densities and unit weights were recorded for all treatments. Significant treatment effects on the rate processes were largely confined to the 21-d period immediately after weekly cutting of rested swards had resumed. On previously rested compared with weekly cut swards, clover leaf appearance rates were increased by 40% and branching rates by 164%. During the same period, grass leaf appearance rates were reduced by 50% and net tillering changed from positive to negative values. Though the rate responses were transient, effects were still apparent at harvest in September, when population density and content (proportion by population density and weight) of clover were significantly higher in the late rest treatment. The variety Kent showed a consistent, though usually nonsignificant, higher leaf appearance and branching rate compared with Milkanova, and in September was characterized by a higher population (7400 m^?2 compared to 3200 m^?2) of smaller units (27 compared to 46 mg/apical meristem) than Milkanova. The results are discussed in relation to defoliation effects and the role of light quantity and quality as they influence the component growth processes. Attention is drawn to the importance of canopy structure and the climatic and/or phenological differences in the relative seasonal behaviour of clover and grass, together with varietal variation within species in influencing responses to management manipulations.