Effects of Competition upon the Productivity of White Clover-Perennial Ryegrass Mixtures. Genetic Analysis

A diallel cross amongst four contrasting white clover varieties was used to provide genetic information on those characters which affect the productivity of white clover-perennial ryegrass mixtures when grown under a cutting-only management regime. Generally speaking those characters recorded on the clover families and in mixtures exhibited highly significant additive and non-ad dative genetic effects, though for1 sortie characters the latter could not be ascribed solely to dominance effects. For two of the clover characters, stolon and leaf number, the genetic picture was radically altered in mixtures by the effects of inter-specific competition Pseudo-genetic effects were also detected for the grass characters, bur it was suggested that the two ryegrass varieties used here were exhibiting phenotypic plasticity. As a group the four intra-varietal crosses performed significantly better than the six intra-varietal crosses for all clover and mixture characters. Only one of the inter-varietal crosses had significantly positive specific combining ability effects for most characters and that was between the two long established varieties ‘S. 184’ and ‘S. 100’. The implications of these results for the breeding of white clover varieties for use in mixtures are briefly discussed.     

Effects of Competition upon the Productivity of White Clover-Perennial Ryegrass Mixtures. Seasonal Trends

Seasonal trends among those characters, affecting productivity in white clover-perennial ryegrass mixtures were investigated m a set of 20 mixtures using clover material derived from a. diallel cross between four contrasting white clover varieties. The productivity and associated vegetative characters showed the familiar pattern of high grass production in spring, and early summer, with a secondary peak of growth in autumn, whilst clover achieved its; highest productivity levels in mid-summer. A significant linear relationship was established between clover canopy height and ryegrass canopy height for all 20 mixtures. Although no significant heterogeneity was detected among the fitted regression lines, there was an indication that the linear regressions provided a better fit to the data in those mixtures having ‘Skomer’ as the grass companion. An analysis of the differences in canopy height between ryegrass and clover indicated that the ‘Menna ‘/’S. 23’ mixture had a reduced seasonal range compared to the other mixtures. Consequently, during the early par', of the growing season, when the tallest canopies occurred and. when shading of clover by the grass was most intense, the difference in canopy height between ‘S. 23’ and ‘Menna’ was relatively low. There was a pronounced effect of grass companion upon canopy height difference, due entirely to the taller growth habit of the ‘Skomer’ ryegrass variety, These interrelationships are discussed briefly in the context of improving the productivity of white clover-perennial ryegrass mixtures.     

Contribution of White Clover Varieties in High-Productivity Systems under Grazing and Cutting

Two medium leaf size varieties of white clover (Trifolium repens L.) were grown in mixture with a range of grass species and varieties in a field experiment conducted over 4 years. The clovers were grown singly or as a blend in swards with a tetraploid perennial ryegrass (Lolium perenne L.), a diploid Italian ryegrass (Lolium multiflorum L.) or tetraploid hybrids between these two species (Lolium x boucheanum Kunth). These grasses were also grown in monoculture. Swards were managed in a regime involving two periods of sheep grazing, two periods of cattle grazing and a silage cut in the same year for 4 years. The objectives of this experiment were 3‐fold: (1) to examine the yield and persistency of white clover in highly productive systems with modern perennial, hybrid and Italian ryegrasses; (2) to compare the performance of white clover varieties grown singly in mixture with grasses with their performance together as a blend, and (3) to examine the persistency of new Italian ryegrass varieties, exemplified by AberComo. The clover content of the sward remained high throughout the experiment, with a mean of 31 % overall. Differences in clover yield between plots were largely attributable to the effects of companion grasses. Total yields were very similar in plots containing different clover varieties and in plots where these varieties had been grown singly or as a blend. Grass yields in mixture were greater than those in monoculture but showed similar relative rankings. The diploid Italian ryegrass variety AberComo showed an unexpectedly high yield in the 4th year of the experiment. The implications of these results for strategies for productive, sustainable agriculture using modern white clover and ryegrass varieties are discussed.     

Assessment of Cattle Preference for Perennial Ryegrass Varieties in Association with White Clover. Does White Clover Content Interfere?

There is a need to evaluate grass varieties under grazing management to allow selection of varieties to increase animal performance. However, the evaluation of animal performance under grazing with each variety is arduous and time-consuming. An alternative is to describe the preference by grazing animals when given a choice of different varieties. Indeed, there is some indications that preference is related to variety characteristics such as leaf content or digestibility, characteristics having also an impact on the level of intake of this variety. However, preference for grass varieties, when sown alone, may not reflect agricultural practice as grasses often occur in mixed swards, associated with white clover. So, the aim of the present work was to determine whether, under grazing management, cattle expressed selections when given a choice of mixed swards in which different perennial ryegrass varieties were grown in association with a common white clover variety at two levels of fertilizer application (60 or 160 kg N ha⁻¹ annum⁻¹). The preference for perennial ryegrass variety per se was then assessed after taking white clover content into consideration. The results show that there were significant differences in preference for perennial ryegrass varieties, per se, even for varieties with the same heading date and ploidy. The effect of white clover cover on sward selection was more marked in swards which received the higher level of nitrogen fertilizer application which may have been related to a more variable white clover content between swards. This suggests that, under low nitrogen fertilizer application, differences in perennial ryegrass varieties preference can be determined in the same experiments as are used to determine ryegrass varieties persistence in association to white clover. Nevertheless, before such ‘variety preference’ tests can be applied in practice, a validation of the possible correlation between ‘variety preference’ and ‘variety intake’ will be necessary.     

Interference Effects in Grass Varieties Grown as Pure Stand, Complex Mixture and Binary Mixture with White Clover

The study aimed at: i) assessing the agronomic value of the pure stand (PS), of the complex mixture (CM) and of the binary mixture with each of six white clover genotypes (MWC) of eight grass varieties belonging to four species and contrasting for earliness of flowering; and ii) investigating the relationships of competitive abilities in CM and MWC with some morpho-physiological traits of the grass varieties. CM showed a 24 % dry matter yield (DMY) advantage over mean value of PS but did not outyield the pure stand of the best yielding component. For grass-white clover associations, the DMY advantage as Relative Yield Total over PS of the components averaged 43 %. DMY of the grass varieties averaged across MWC conditions was related positively to total DMY (r = 0.99) and negatively to associated clover DMY (r = - 0.96). Competitive abilities as Relative Yield of the grass varieties in CM and in MWC correlated (r = 0.84) and were both positively associated to DMY, canopy height and inclination of leaves in PS. The results suggested that association of white clover with a grass of moderately high vigour, intermediate earliness and relatively erect leaves may provide the best option for rotational meadows in the experiment environment.     

Quality characteristics of backcross hybrids between Trifolium repens and Trifolium ambiguum

Two generations of novel backcross hybrids between white clover (Trifolium repens L.) and the related species Trifolium ambiguum,with white clover as the recurrent parent, were analysed for forage quality characteristics. Water soluble carbohydrate (WSC) levels, nitrogen concentrations (%N), and dry matter digestibility (DMD) were recorded for the hybrids and parental species grown in deep soil bins with and without a companion grass. Half the experimental material was subjected to moisture stress by being left unwatered through the four-week period when measurements were made. Four such four -week cycles of drought were imposed. The first and second generation backcrosses had the highest values for WSC, but were lowest for N%. There was no difference between the different legume lines (parents and backcrosses) for DMD. No effects of either moisture stress or presence of a grass companion on quality traits were observed. The WSC and %N of the grass component differed significantly depending on which legume line it was grown with. The importance of these results for the potential use of these hybrids in agriculture is discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Breeding white clover for tolerance to low temperature and grazing stress

Summary Low temperature and grazing are the two major stresses limiting white clover yield in mixed swards grown in the marginal areas of the UK. White clover has traditionally been used to improve productivity in such areas but is increasingly being used to reduce input costs in more productive areas. Considerable genetic variation exists in characters associated with cold tolerance, low temperature growth and grazing tolerance. This paper describes recent progress in developing varieties tolerant to three major stresses and which give greater and more reliable production.     

生草栽培对柑桔丛枝菌根形成及果实品质的影响

试验以枳砧锦橙(CitrussinensisOsbeck.)初结果树为试材。试验设4个处理,分别为柑桔园间作百喜草(Bahiagrass)(PaspalumnotaturnFlugge)、白三叶草(Whiteclover)(TrifoliumrepensL.)、紫花苜蓿(Alfalfa)(MedicagosativaL.)和多年生黑麦草(Perennialryegrass)(LoliumperenneL.),以清耕为对照。单行小区,随机区组排列,3次重复。结果表明,在干旱季节,所有生草处理区都显著改善了根际土壤水分和温度状况,而百喜草和白三叶草生草区土壤丛枝菌根真菌孢子数和柑桔根系菌根侵染率显著高于清耕区和其它生草区。表明在干旱季节,百喜草和白三叶草生草栽培促进了果园土壤丛枝菌根真菌的生长和柑桔根系菌根的形成,从而促进了柑桔根系对水分和磷素营养的吸收利用。试验结果表明果园管理中宜选择菌根侵染率高的草种如百喜草、白三叶草在柑桔园进行生草栽培,促进柑桔根系菌根形成,改善植株生理代谢,提高果品质量。     

Selection of timothy (Phleum pratense L.) under different conditions of culture

Summary Recurrent selection for herbage weight under different conditions of competition and cutting frequency between hexaploid timothy cultivars and within their open-pollinated progenies was inefficient in producing in two generations either highly improved material as compared to the control (Climax) or any pronounced cultural ecotypes as shown by genotype x treatment interaction analyses, but some Climax and Hungarian selections were promising.Parent-offspring correlations indicated high heritability of heading and morphological characters in the first generation but not in vigor and yield which were the only characters measured in the second generation. The correlations may have been enhanced by the technique of open-pollination. A few selfings included showed very high inbreeding depression.Very early characters were useful in indicating growth types. Plant x season interactions were obvious but not tested. There were grass x legume interactions in the F₁ showing the different effects of alfalfa and white clover. A leaf marking was useful in differentiating white clover cultivars. In the F₂, spacing x cutting system interactions were significant. Light interception was significantly correlated to yields in the grass rows. Discriminant functions indicated the importance of tiller thickness for a selection index.The results are discussed in relation to experimental errors and to the possible polysomic segregation in hexaploid timothy.Published with the approval of the Director of the Ottawa Research Station, Central Experimental Farm, Ottawa. Contribution No 264.     

Symbiotic Nitrogen Fixation of White Clover in a Mixed Sward is not Limited by Height of Repeated Cutting

The aim of this study was to investigate whether symbiotic nitrogen fixation in white clover nodules limits nitrogen supply and hence clover growth by repeated defoliation at two cutting heights. Other possible factors governing symbiotic nitrogen fixation in the field were also elucidated. Using ¹⁵N, a 2-year field experiment including white clover ( Trifolium repens L. cv. Ladino) and perennial ryegrass ( Loliumperenne L. cv. Bastion) in monocultures and in mixtures was conducted in Eschikon, Switzerland. The effect of two cutting heights (4 cm and 10 cm above ground level) on the performance of symbiotic nitrogen fixation of white clover in the different sward-types was investigated. After each harvest, the plots were fertilized with 3 g N m^-2(equivalent to 30 kg N ha^-1 cut^-1 or 210 kg N ha^-1 year^-1). In both years, white clover grown in a mixture with grass received a significantly higher percentage of nitrogen from symbiotic fixation compared with clover grown in monoculture. This phenomenon is attributed to the strong competitiveness of ryegrass in soil nitrogen uptake. Consequently, white clover in the clover-ryegrass mixture was more dependent on symbiotic nitrogen fixation than that grown in monoculture. The cutting height did not preferentially influence symbiotic nitrogen fixation, as opposed to the uptake of mineral nitrogen from the soil. From this finding it is suggested that symbiotic nitrogen fixation did not limit the supply of nitrogen to clover and hence its growth. It is proposed that symbiotic nitrogen fixation in white clover is regulated by the demand for nitrogen rather than by the availability of carbohydrate reserves in the stolons. Symbiotic nitrogen fixation should thus be looked upon as an integrated plant growth factor and not as an isolated phenomenon.     

白三叶种质资源主要数量性状的变异与相关性研究

为了探讨白三叶(Trifolium repens L.)种质资源农艺性状变异和相关性,揭示各种质材料的特征特性和种质资源群间的遗传关系,本研究对78份白三叶种质的12个主要数量性状的变异和相关性进行了分析。结果表明：这些种质材料存在着丰富的遗传多样性,各个数量性状间均呈正相关,植株高度与小叶长度、小叶宽度、小叶大小、叶柄长度呈显著正相关;小叶长度与小叶宽度、小叶大小呈极显著正相关,与叶柄长度、叶柄直径呈显著正相关;小叶宽度与小叶大小呈极显著正相关,与叶柄长度、花柄直径呈显著正相关;小叶大小与叶柄长度、叶柄直径、花柄直径呈显著正相关。该批种质材料能为各类白三叶育种和遗传研究提供较为丰富的亲本材料。     

河南省优良草种区划的探索

通过对河南省主要草种的生物学特点、生态学特点以及经济价值的分析,结合当前农牧业发展对饲草的需要特点的分析,采用系统学优化理论,确定了河南省当家草种应具备的条件,制定了全省草种区划方案。结果表明：适合河南省的当家草种有10种,分别是：紫花苜蓿、沙打旺、苇状羊茅、多年生黑麦草、小冠花、红三叶、杂三叶、鸡脚草、无芒雀麦;适合河南省草业优质发展的草种区域有6个,分别是：豫北平原牧草种植区、豫西北黄土丘陵和太行山地种植区、豫中东低洼平原和南阳盆地种植区、豫西山地种植区、淮南与桐柏山区种植区、豫东平原种植区,并且明确了各区域内以当家草种为核心的草种优化组合。河南省当家草种的制定和草种区划的确定有利于为河南省各地草种的正确选择和草业未来大发展提供理论依据。     

白三叶基因工程改良研究进展

白三叶（Trifolium repens L.）作为优良牧草和地被植物具有重要的应用价值。但其耐盐碱、干旱能力差等原因致使应用受到广泛限制,转基因技术的快速发展为白三叶种质创新提供了有效技术手段,为开展白三叶分子育种奠定了基础。本文就近30年来有关白三叶组织培养、转化方法、转基因遗传改良方面的研究进展进行了综述,并对其应用进行了展望。     

Interspecific hybridization in the genus Trifolium

The genus Trifolium (Leguminosae) or clovers includes a number of agriculturally important forage species. In particular Trifolium repens (white clover) and Trifolium pratense (red clover) are widely used in temperate livestock agriculture. Interspecific crossing programmes within this genus have been carried out for more than 50 years, in many cases involving embryo rescue or ovule culture techniques. In general, the aims of these have been twofold: to aid understanding of evolutionary relationships within the genus and to introgress useful traits, predominantly into white or red clover. In the case of white clover, the greatest efforts have been made with respect to crossing with Trifolium nigrescens and Trifolium ambiguum . Novel germplasm has been developed and useful traits transferred, although no material has yet been commercialized. The main emphasis with respect to red clover has been increasing longevity through crosses with more persistent species, particularly Trifolium medium . Again, this work has not yet resulted in new varieties. Wider crossing programmes, allied in recent years to comparisons at the DNA level, have shed considerable light on the systematics and evolution of the genus. The future of interspecific hybridization for improvement of the agriculturally important clovers and the role of molecular approaches in this are discussed.     

Effects of Interspecific Competition on the Inheritance of Quantitative Traits in White Clover *)

Analysis of a half diallel cross among four white clover cultivars, grown as monocultures and in mixtures (duocultures) with two separate perennial rye-grass cultivars, confirmed that for stolon and leaf number a higher proportion of the genetic variance in duocultures was non-additive. For canopy height and dry weight, however, this proportion was less affected by inter-specific competition. An additive-dominance model of gene action sufficed for all four characters in monocultures, but in duocultures epis-tasis was just one of the factors complicating the genetic picture. This, coupled with short-term reversals in the direction of dominance, merely illustrates the complexity of breeding white clover cultivars. Possible future developments in the breeding of white clover are discussed in the light of these results.     

Nitrogen Fixation in Hybrids of White Clover (Trifolium repens L.) and Caucasian Clover (Trifolium ambiguum M. Bieb)

Hybrids between white clover (Trifolium repens L.) and Caucasian clover (or Kura clover, Trifolium ambiguum M. Bieb) are a potential route for the improvement of drought tolerance and persistence in white clover. However, to be agronomically viable they must show no significant reduction in their potential for nitrogen fixation relative to white clover. A comparative study of growth rate and nitrogen fixation was carried out in flowing solution culture without a supply of mineral nitrogen to the plants. The two parental species and two generations of backcross hybrids, with white clover as the recurrent parent, were assessed. The growth rate and N content of T. ambiguum were significantly lower than those of the other lines. However, dry matter production, nodule biomass per plant and rates of fixation were similar in second‐generation backcross plants and white clover. The results suggest that the agronomic potential of this novel germplasm is not compromised by limitations with respect to nitrogen fixation.     

Introgression of reproductive traits from Trifolium nigrescens increases the seed yield of white clover (T. repens)

Interspecific hybrids between white clover and the annual, profuse flowering ball clover have been developed as a strategy to improve the potential seed yield of white clover. The seed yield and seed yield components of advanced white clover × ball clover hybrids were compared with white clover in replicated field experiments over two harvest years. The backcross 3 hybrid had more reproductive nodes per stolon than the control variety ‘AberHerald’ which was reflected in more inflorescences per unit area but inflorescences with fewer florets and seeds. The hybrid had a greater potential seed yield than the control variety in 2002 but not 2001. Backcross 3 hybrids have also been produced across the range of leaf size categories. Medium and large leaved hybrids produced significantly more inflorescences and a higher potential seed yield than control varieties of comparable leaf size in both harvest years. The small leaved hybrid was not significantly greater than the control variety ‘AberDale’ in either inflorescence production or seed yield.     

The characteristics of European,Mediterranean and other populations of white clover (Trifolium repens L.)

Indigenous populations of white clover (Trifolium repens L) collected from Europe, the Mediterranean area, as well as other parts of the world have been grown as spaced plants at Aberystwyth. Data were collected on various morphological characters, green weight and leaf marks.Leaflet length, width, area, plant height and green weight were all strongly correlated, but not the flowering date or leaflet shape (= ratio length/width).The mean leaflet size of Mediterranean populations (the supposed centre of origin) was very variable, and decreased northwards. All British populations were uniformly small leaved.Most of the populations were much less vigorous than the bred control varieties and only a few equalled them. Nevertheless some of the introductions exhibited seasonal growth patterns, especially growth into the autumn, and large leaves that could be usefully incorporated in improved varieties. Populations originating one step back towards the centre of origin will probably be the most useful when immediate adaptation is the aim.     

Heritability of, and genetic correlations among, forage and seed yield traits in Ladino white clover

Sixteen white clover genotypes and their half-sib progenies were grown in pure stand. Each clover progeny was also grown in a mixed stand that also included three grass varieties of different species. Dry matter (DM) yield was measured over 2 years and seed yield at the second year. Competitive ability of clover families was defined as the ratio between mixed stand and pure stand for clover DM yield. Narrow-sense heritability from parent-offspring regression of pure stand data was high to moderate (h²0.50) for DM yield, seed yield and most of their components. Persistence as predicted by stolon density showed negative genetic correlations with seed yield (r_g= -0.70) and DM yield (r₂= -0.60), whereas seed yield and DM yield tended towards a negative correlation (r₂=0.45). Sizes of different vegetative or reproductive organs were generally correlated positively. Evidence was provided that the set of parent genotypes represents well the genetic variation available within the Ladino gene pool. The results highlighted the difficulty of combining relatively short-term DM yield, persistence and seed yield into a unique plant type. Differences in competitive ability emerged only in the second year, better ability being related mainly to petiole length among the traits assessed in pure stand.     

Self-compatibility and heterosis in white clover (Trifolium repens L.)

Inbred lines of white clover (Trifolium repens L.) have been produced utilising the rare self-fertility (Sf) allele. Twenty-two lines of four distinct groups have been maintained through five generations of selfing by single seed descent. Fourteen lines were used in crosses to produce F¹s. Both parents and hybrids were analysed for a range of morphological characters. Positive heterosis for dry matter production was observed in half the hybrids. No other trait showed significant heterosis. The degree of heterosis appears to be related to the extent of variation in morphological characters between the parental lines, some combinations of lines yielding heterotic hybrids in all cases, others in none. The majority of the F¹ hybrids are superior in terms of dry matter production to eight control varieties. This revised version was published online in July 2006 with corrections to the Cover Date.