The effects of frequency of cutting and cultivar on the period of leaf expansion in white clover grown in mixed swards

The period of leaf expansion in white clover ( Trifolium repens ) grown with perennial ryegrass ( Lolium perenne ) in mixed swards and the relationship between the period of leaf expansion and leaf appearance rate were examined in a field experiment in The Netherlands in which two cutting frequencies (low and high) and three white clover cultivars (Alice, Retor and Gwenda) were used. A significant positive correlation existed between the period of leaf expansion and leaf appearance rate under a low frequency of cutting, indicating that a longer cutting interval might allow the expression of cultivar and environmental differences in both leaf expansion and leaf appearance rate. In most cases, frequent cutting tended to shorten the period of leaf expansion, which may be an important cause of lower dry-matter yield in the high frequency of cutting treatment. The large-leaved cv. Alice had a significantly longer period of leaf expansion than the other cultivars under frequent cutting in autumn. This might partly account for the mixtures with Alice having a greater proportion of white clover than the mixtures with Retor and Gwenda in autumn. Some of the variation between low- and high-frequency cutting treatments in the period of leaf expansion could be explained by mean and minimum temperatures (38% and 15% respectively).     

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.     

Severity of damage to Trifolium repens leaves by certain invertebrate species in mixed perennial ryegrass/white clover swards: response to cultivar, cutting frequency and sward characteristics

The severity of damage by certain invertebrate species to white clover (Trifolium repens) leaves on the main stolons of plants grown in mixed perennial ryegrass/white clover swards was examined in a field experiment in the Netherlands in which two cutting frequencies (high and low) and three white clover cultivars (Retor, Alice and Gwenda) were used. The damage to the leaves was described in terms of the numbers of damaged leaves and the extent of that damage (slight <20%, moderate 20–50% and heavy >50%). The relationships between leaf damage and sward characteristics (white clover content, above-ground biomass and sward height) were evaluated throughout the growing season. Over the whole experimental period, 23·7% and 27·4% of the total number of leaves produced per stolon were damaged by slugs and weevils in the low- and high-frequency cutting treatments respectively. High-frequency cutting increased the number of leaves in the total leaf damage and moderate leaf damage categories by 21·4% and 34·8%, respectively, compared with the low-frequency cutting. The cv. Retor (medium-leaved) experienced the most severe damage by invertebrates. It had much higher leaf damage than cvs Alice (large-leaved) and Gwenda (small-leaved) at either cutting frequency, both in the total number of damaged leaves and in the different damage categories. Differences among cultivars in the number of damaged leaves and relative leaf damage occurred primarily in spring, late summer and autumn, but did not differ during the early- and mid-summer months. This study indicates that variations in leaf damage among clover cultivars were associated with differences in measured sward characteristics. Both the number of damaged leaves and the relative leaf damage were strongly negatively correlated with white clover content and biomass in spring, late summer and autumn under each cutting treatment. White clover content and biomass explained 65%, 59% and 50% of the variation in the number of damaged leaves in spring, late summer and autumn, respectively, and 58%, 57% and 45% of the variation in relative leaf damage in these three periods. Thus, sward characteristics may play a role in regulating the severity of invertebrate damage to clover leaves in addition to the primary effects of HCN.     

Competition in perennial ryegrass–white clover mixtures under cutting. 1. Dry-matter yield, species composition and nitrogen fixation

Persistence of white clover (Trifolium repens) in mixtures was studied in a long-term experiment. Mixtures of two cultivars of perennial ryegrass (Lolium perenne) with contrasting growth habits and three white clover cultivars differing in leaf size were sown in 1991 and evaluated at two cutting frequencies. During 1995 and 1996 mixtures with large-leaved white clover cv. Alice had the highest dry-matter (DM) content, clover and N yield, and the highest white clover content, and mixtures with medium-leaved Retor the lowest, whereas mixtures with small-leaved Gwenda yielded most grass DM. In 1995 averaged over cutting treatments and mixtures, the mixtures yielded 11·8 t DM ha^?1 with a white clover content of 0·6; the apparent N fixation was 393 kg N ha^?1. In 1996 these values declined to 8·5 t DM ha^?1, 0·48 white clover and 236 kg N ha^?1. There was no significant effect of cutting frequency on DM yield or white clover content, whereas the effects of grass cultivar were not consistent. In spring there was a peak in the DM production of the mixtures, coinciding with a peak in production of the grass component. However, in summer and autumn the seasonal pattern of DM production of the mixtures was similar to that of the white clover component. Both cultivars of perennial ryegrass showed the same seasonal response, but the seasonal growth pattern of white clover differed slightly between clover cultivars and cutting treatments. In later years only one cutting frequency was imposed, and no yield measurements were taken. White clover was judged to have performed well during 1997; the clover content in September was very high (0·76), whereas in October 1998 it was 0·45. Mixtures with Alice contained most white clover. Despite fluctuations in white clover content during 1991–98, all clover cultivars had persisted 7 years after sowing, irrespective of companion grass cultivar, at both cutting treatments.     

Effects of white clover cultivar and companion grass on winter survival of seedlings in autumn-sown swards

The aim was to study the effects of white clover cultivar and combinations with perennial ryegrass cultivars on seedling establishment in autumn‐sown swards and on winter survival of seedlings. Large‐leaved white clover cv. Alice and small‐leaved white clover cv. Gwenda, and an erect and a prostrate perennial ryegrass cultivar were sown in autumn in pure stands and as four binary grass‐clover mixtures. Mixtures of white clover cv. Huia and Aberherald with perennial ryegrass were also sown. Companion grasses had no significant impact on the establishment of white clover. The number of seedlings of white clover cv. Alice in mixtures (335 m^?2) was higher than cv. Gwenda (183 m^?2) and pure swards had similar white clover population densities as mixed swards. White clover cv. Huia tended to have more seedlings than Aberherald (355 and 205 m^?2 respectively). No stolons were produced prior to a severe winter, because of the late sowing date. Winter survival of clover seedlings was 0·56 in mixtures and 0·69 in pure stands, irrespective of white clover or companion grass cultivar. Stolon development of white clover in autumn is often considered essential for overwintering survival and spring growth. In this study, there was considerable survival of the non‐stoloniferous tap‐rooted seedlings of all four clover cultivars despite a severe winter.     

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.     

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.     

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.     

White clover seed production from mixed swards: effect of sheep grazing on stolon density and on seed yield components of two contrasting white clover varieties

A trial was carried out over two harvest years to assess the effect of sheep grazing and closing date on stolon density and seed yield components of two contrasting white clover varieties and to determine the potential for producing clover seed from mixed swards in an integrated livestock/seed production system.
Overall, the small-leaved cv. S184 had a higher stolon density at closing and harvest than the large-leaved cv. Olwen, but only significantly so in 1987. Stolon density at closing was generally increased by delaying closing. Varieties also differed in their response to closing date. Stolon density of cv. S184 in both harvest years, generally increased with later closing whilst cv. Olwen was less influenced by closing date and at all dates in 1987, and all but one date in 1986 was not significantly different from the ungrazed treatment.
Delay of closing significantly increased stolon density at harvest on all closing dates in 1987 but had no effect in 1986. Varieties responded similarly to a delay in closing but the magnitude of increase of cv. Olwen was less than that of cv. S184.
The seed yield components of both cv. S184 and cv. Olwen were influenced by closing date, but the effect differed between years. Although there was some initial increase in inflorescence production, delaying closing after inflorescence buds appeared on the stolon reduced inflorescence number of cv. Olwen in both years but reduced inflorescence number in cv. S184 in 1986 only. However, inflorescence size, seed yield per inflorescence and the proportion of ripe inflorescences at harvest was reduced in both years. Cultivar S184 produced more inflorescences than cv. Olwen in both years and in both years tolerated later closing than cv. Olwen.
The effect of sheep grazing and closing date is discussed in relation to stolon removal and the suitability of particular leaf types for this system of seed production.     

White clover growth patterns during the grazing season in a rotationally grazed dairy pasture in New York

White clover (Trifolium repens L.) is an important stoloniferous pasture legume in the Great Lakes region of the United States, but it often has limited persistence. Researchers in New Zealand and Wales have found that in spring, compared with other seasons, white clover plants have reduced branching complexity and have the fewest buds that produce leaves. They therefore suggested that in spring the plants are most vulnerable to grazing and climatic stress. Because of severe winter and cool, wet spring weather in New York State, it was hypothesized that white clover plants would also be of low branching complexity, smaller and have low axillary bud activity in spring compared with later in the grazing season. To test this, growth of white clover was monitored in an orchard grass (Dactylis glomerata L.)/white clover pasture in New York that was rotationally grazed with dairy cows during the 1993 and 1994 grazing seasons. Three sets of plants were sampled. The first set consisted of forty random plants sampled before each grazing event. Stolon branching order, number of each stolon branching type and area the plant occupied were determined. Approximately each month before one grazing event, a separate set of 32 random plants was measured in the field to determine the area they occupied; these plants were then removed to the laboratory for the measurement of stolon order, number of each stolon type, stolon lengths, total number of growing points, number of taproots and adventitious roots, root position and above-ground dry matter. Once a month, 12 additional plants were removed to measure axillary bud activity at each node. Leaf development from nodes tended to increase from spring to summer. However, the stolon branching order of white clover plants was not simpler in spring compared with summer or autumn. In 1994 during and after a dry and hot period, white clover plants were smaller, of lower stolon branching order and had fewer roots. Climate and associated soil organism activity appear to explain the different white clover growth patterns observed in New York and New Zealand. Severe winters in New York limit earthworm activity and stolon burial, which is important in contributing to stolon/plant breakdown in New Zealand. During the years of this study in New York, a hot and dry period had the most negative effect on the growth pattern of white clover.     

The effect of stolon burial and defoliation early in the growing season on white clover performance

A factorial pot-experiment was carried out to investigate the effect of burial of stolons (stolons unburied or buried to a depth of 0·5–1 cm at day 0), with and without defoliation (plants uncut or all unfolded laminae removed at weekly intervals) on the growth of two varieties of white clover (cv. Kent or cv. Milkanova) harvested on three dates (14 d, 28 d and 42 d after burial). The soil used was a 3:2:1 mixture of clay loam:peat:sand, and there were three replicates of each variety for each treatment at each harvest date. Burial of stolons in the absence of defoliation had no effect on stolon extension, leaf appearance, or the concentration of water-soluble carbohydrate in the stolons. The number of axillary buds developing (new branches plus flowers) was increased on the new surface-growing tips of primary stolons but was reduced on branch (i.e. secondary) stolons. The proportion of branches to flowers was largely unaffected by burial. Defoliation caused substantial reductions in the concentration of water-soluble carbohydrates in stolons and stolon extension growth, a reduction in number of axillary buds developing, and in the proportion of buds which were floral, but had only a small effect in reducing leaf appearance. The combination of stolon burial and defoliation resulted in the death of secondary stolons; 42% of all secondary stolons had died by day 42, and stolon extension, leaf appearance and numbers of axillary buds developing on secondary stolons were severely reduced. The numbers of axillary buds developing on primary stolons was increased owing to more buds developing on the resurfaced stolon tips, but the increase was inadequate to compensate for the reduced growth, and death of secondary stolons. Differences between clover varieties in response to treatments were small. In general, the smaller variety, cv. Kent, was more adversely affected by burial than the larger variety, cv. Milkanova. A variety x defoliation interaction occurred for senescence of leaves; petiole senescence of cut leaves was reduced or similar to that of intact leaves for cv. Kent, but was increased for cv. Milkanova. The results are discussed in relation to treatment effects on carbon sources and sinks, and in relation to climate-soil interactions.     

Factors affecting the stolon growth of white clover in ryegrass/clover patches

The effects of spatial location of white clover ( Trifolium repens L.) within a perennial ryegrass ( Lolium perenne L.)/white clover pasture on stolon and petiole extension were investigated in two experiments, where patch size containing white clover (0·5 m, 1·5 m and 4 m diameter), location within the patch (inside and edge) and cutting height (4 cm and 8 cm) were varied. Stolon extension rate was greater on the edge of a patch (12·1 mm week⁻¹) than inside the patch (7·2 mm week⁻¹). Patch size affected both stolon and petiole extension rate, which were both greater in small and medium-sized than in large patches. It is suggested that the fastest spread of white clover in patchy sward environments should occur from small patches, which could double in diameter during a growing season. Manipulating the heights of vegetation within and outside large patches affected light quality (red-far red; R/FR) at ground level, which was greater under shorter than taller swards and greater under the canopy of the grass matrix than the grass/white clover patch. However, the height differences between adjacent vegetation had little effect on stolon or petiole growth. In May only, stolon extension at the patch boundary was greatest when both patches and the grass matrix had a height of 8 cm.     

Performance of white clover/perennial ryegrass mixtures under cutting

Clover persistence in mixtures of two varieties of perennial ryegrass (Lolium perenne) with contrasting growth habits and three white clover (Trifolium repens) varieties differing in leaf sizes was evaluated at two cutting frequencies. An experiment was sown in 1991 on a clay soil. The plots received no nitrogen fertilizer. In 1992, 1993 and 1994, mixtures containing the large-leaved clover cv. Alice yielded significantly more herbage dry matter (DM) and had a higher clover content than mixtures containing cvs Gwenda and Retor. Companion grass variety did not consistently affect yield or botanical composition. Cutting at 2 t DM ha^?1 resulted in slightly higher total annual yields than cutting at 1.2 t DM ha^?1, but did not affect clover content. In 1992 the mixtures yielded, depending on cutting frequency and variety, 10·6–14·6 t DM ha^?1 and 446–599 kg ha^?1 N, whereas grass monocultures yielded only 1·2–2·0 t DM ha^?1 and 25–46 kg ha^?1 N. From 1992 to 1994 the annual mean total herbage yield of DM in the mixtures declined from 12·2 to 10·5 to 8·7 t ha^?1, the white clover yield declined from 8·7 to 6·5 to 4·1 t ha^?1 and the average clover content during the growing season declined from 71% to 61% to 46%, whereas the grass yield increased from 3·4 to 4·0 to 4·5 t ha^?1. The N yield decreased from 507 to 406 to 265 kg N ha^?1 and the apparent N fixation from 470 to 380 to 238 kg N ha^?1. Nitrate leaching losses during the winters of 1992–93 and 1994–95 were highest under mixtures with cv. Alice, but did not exceed 10 kg N ha^?1. The in vitro digestible organic matter (IVDOM) was generally higher in clover than in grass, particularly in the summer months. No differences in IVDOM were found among clover or grass varieties. The experiment will be continued to study clover persistence and the mechanisms that affect the grass/clover balance.     

Effects of shoot,root and stolon temperature on the development of the potato (Solanum tuberosum L.) plant. II. Development of stolons

Summary Cultivars Bintje and Désirée were grown in a multi-compartment set-up, which allowed different shoot, root and stolon temperatures, to study the effects of temperature around different plant organs on the development of stolons. A high root temperature induced orthotropic stolon growth in cv. Bintje. Stolon number was not affected when temperature was increased in one compartment only, but high root temperature combined with high stolon temperature or high air temperature reduced stolon number. For ‘Bintje’, branching of stolons and their dry-matter yield were enhanced by a high temperature in each compartment, but combining high air temperature with another temperature increase was detrimental. Stolon initiation was less synchronized when plants were exposed to both high air temperature and high stolon temperature. The hormonal regulation of stolon formation and the effects of temperature on the hormonal balances involved are discussed.     

The effect of nitrogen in spring on shoot number and leaf area of white clover in mixtures

Swards of perennial ryegrass/white clover were cut to 3 cm in March and allowed to regrow for approximately thirteen weeks. Half the swards received nitrogenous fertilizer at the time of cutting (+N treatment) while the other half received none (—N treatment).
About four weeks after the application of treatments, the number of clover shoots in the — N treatment was significantly higher than that in the +N; at this stage total leaf area index on both treatments was low, but significantly greater in the + N. After 30 days, more light was reaching ground level in the — N treatments, and about 40 days after cutting there was between 17 and 50% more clover in the — N treatment than the +N.
It is suggested that the inhibition of stolon branching with a consequent restriction in the rate of increase of clover leaf area is a major factor in the adverse effect of N fertilizer on clover.     

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.     

The effect of row spacing and cover-crop on stolon development and the seed yield components of white clover cultivars of contrasting leaf size

Three white clover cultivars, S184 (small-leaved), Menna (medium-leaved) and Olwen (large-leaved), were broadcast or sown in 15, 30 or 60 cm drills at a seed rate of 3 kg ha^-1. The three cultivars were either sown without a cover-crop or sown under a cover-crop of spring barley (cv. Crescent) or peas (cv. Countess). The effects of these methods of establishment on the stolon growth and components of seed yield were subsequently measured.
Stolon growth and development was influenced by row spacing, cultivar and season. The overall plant response at all but the widest row spacing (60 cm) was to increase stolon growth such that inflorescence production, the number and proportion of ripe inflorescences and the other seed yield components were unaffected by row spacing. Cultivars differed in their response to row spacing. Cultivar Olwen produced most inflorescences and more ripe inflorescences when broadcast, cv. S184 when sown at 60 cm row spacing and cv. Menna at 15 or 30 cm row spacing. Cultivars also differed in their response to cover-crop, with cvs Menna and S184 producing more inflorescences and more ripe inflorescences when sown under barley and peas than when pure sown. The inflorescence production of cv. Olwen was not influenced by cover-crop.
The relationship between vegetative and reproductive growth is discussed in relation to establishment, cultivar and climate and the possible implications for the establishment of white clover seed crops in the UK.     

Growth and colonization of a range of white clover cultivars strip-seeded into an upland perennial ryegrass sward

Seven cultivars of white clover (Trifolium repens L.) (Kent, S184, Huia, Menna, Donna, Alice and Nesta) and a commercial mixture, ‘Ensign’, were strip-seeded into an upland perennial rye-grass (Lolium perenne L.) sward in late June 1986. Swards were first grazed by sheep, either on 5/6 August (early) or on 19/20 August (late) and then every 14–21 days (frequently) or 28–42 days (infrequently) during 1986, followed by a common grazing regime in 1987. During April to mid-June 1988 the swards received either a moderate amount of nitrogen or none and were cut frequently or once only in mid-June. Growth of individual seedlings was assessed before and after grazing during 1986 and stolon accumulation and distribution and sward colonization were assessed during 1987 and 1988. All cultivars emerged rapidly and satisfactorily and there were no consistent significant differences in the overall dry matter accumulation per seedling during establishment. During the first autumn the proportion of the aboveground biomass removed during grazing was smallest in Kent (c. 20%) and largest in Nesta (c. 40%). Kent and S184 produced most leaves and stolons and the greatest length of stolons per seedling and per individual stolon, and Nesta and Alice the fewest leaves and stolons and shortest stolons. Seedlings grazed early had heavier and longer stolons than those grazed late; those grazed frequently had more leaves, stolons and growing points than those grazed infrequently, especially following early grazing. During 1987 Kent and S184 had consistently the largest number of stolon growing points, and weight and length of stolons per unit area; these two cultivars and Nesta also colonized the sward more rapidly than the other cultivars. All cultivars contributed substantially and similarly to herbage production in late September. There were no residual effects of the 1986 treatments after the summer of 1987. During 1988 additions of nitrogen fertilizer at 100 kg N ha^-1 or allowing the herbage to remain undefoliated between mid-April and mid-June both independently halved the number of stolon growing points per unit area; together they reduced it by 80%. Nitrogen also, on average, halved stolon weights but less so in Nesta, Alice and Huia and more so in all other cultivars. Infrequent defoliation greatly decreased stolon weights in Kent and S184 but had no significant effects on the other cultivars. Sward colonization was almost complete by June and entirely so by October for all cultivars in all treatments. Implications of the results for the after-management of strip-seeded white clover are discussed.     

The relationships between stolon characteristics, winter survival and annual yields in white clover (Trifolium repens L.)

Five white clover populations of Swiss origin and three bred varieties were grown in binary mixtures with two perennial ryegrass varieties, Aurora and S23. The seasonal yields of clover and grass plus clover were measured under a cutting regime during the second and third years after establishment. A series of destructive detailed sward measurements was made during the late autumn to spring period preceding each harvest year. In this way changes in the amounts of stolon, leaf plus petiole and numbers of growing points were monitored during the winter.
There were large differences in clover yield between populations in both years. These were evident from the first (spring) harvest in each year. Higher-yielding clovers in spring tended to produce higher annual clover yields. No grass × clover interaction was evident at any harvest. Large differences between clovers were also apparent in the morphological characteristics measured, with the Swiss material generally having greater amounts of stolon, leaf plus petiole and numbers of growing points present in early spring. It is proposed that these factors contribute to the high spring yield in the Swiss populations. Loss of stolon length over each winter was less in the Swiss material, indicating that its good spring growth was not obtained at the expense of winter hardiness. Annual clover yield was found to be significantly positively correlated with the amount of stolon present in spring, exemplifying the importance of stolon survival over the winter.     

Evaluation of white clover varieties under grazing and their role in farm systems

Four experiments were established in 1981–84 to investigate the effect of defoliation treatments on white clover varieties when grown with S23 perennial ryegrass. Treatments included a cutting only regime, as used in National List trials, and grazing systems simulating as near as possible those used on farms.
Differential effects of cutting and continuous sheep stocking on white clover varieties, together with significant variety × defoliation interactions, illustrated the importance of the grazing animal in the evaluation of white clover. Differential effects of cattle and sheep grazing were also evident. Cattle grazing was less detrimental to white clover than was sheep grazing, i.e. cattle were not selective, and less stolon was removed. In general, with both cattle and sheep grazing the larger the clover leaf size the greater the loss in stolons, which in turn decreased persistency. The results illustrate how alternating cutting, sheep grazing and cattle grazing managements can be used to maintain optimum clover/grass balance.
Successful clover/grass swards depend on the retention of clover, yet avoiding clover dominance. The difference in N transfer between clover varieties, especially those within the same leaf category, and the extra grass produced without fertilizer N, emphasized the importance of varietal choice. Breeding programmes have been concerned with the selection of larger-leaved, long-petioled varieties for growing in competition with grass in the presence of fertilizer N. However, the present results showed that, under continuous sheep stocking, increase in leaf size does not increase clover yield or persistency.
The results presented emphasize the importance of the grazing animal in the evaluation of white clover varieties and indicate that yield of clover dry matter should not be the major criterion for selection of varieties for farm systems.