Quantitative trait loci for leaf length in perennial ryegrass (Lolium perenne L.)

Leaf length is an important trait for forage grasses. Molecular marker development offers the opportunity to identify Quantitative Trait Loci (QTLs) and to begin to dissect the genetic regulation of complex traits. The objective of this study was to look for QTLs for leaf length and related traits (plant height, lamina and sheath lengths, leaf elongation rate and leaf elongation duration) in the progeny of a cross between two genotypes of Lolium perenne L. with contrasting leaf length. Measurements were performed in a plant nursery in autumn and spring, and in a heated greenhouse in winter. A high level of variability and heritability was observed for all traits. Lamina lengths at different dates were moderately but significantly correlated. For all traits, QTLs were found and explained between 0·08 and 0·44 of phenotypic variance. Although different leaf length QTLs were found at different dates, linkage groups two, four and seven were particularly important.     

Effects of perennial ryegrass (Lolium perenne L.) cultivars on herbage production, nutritional quality and herbage intake of grazing dairy cows

Four perennial ryegrass (Lolium perenne L.) cultivars were compared for differences in herbage production, nutritive value and herbage intake of dry matter (DM) during the summers of 2002 and 2003. Two paddocks were sown with pure stands of four cultivars in a randomized block design with three replicates. Each plot was subdivided into fourteen subplots (22 m × 6 m) which were grazed by one cow during 24 h. Twelve lactating dairy cows were assigned to one cultivar for a period of 2 weeks in a 4 × 4 Latin square experimental design; the experiment lasted 8 weeks in each year. Sward structure (sward surface height, DM yield, green leaf mass, bulk density and tiller density) and morphological characteristics were measured. The ash, neutral‐detergent fibre, acid‐detergent lignin, crude protein and water‐soluble carbohydrate concentrations, and in vitro digestibility of the herbage were measured. The sward was also examined for infestation by crown rust (Puccinia coronata f. sp. lolii). Herbage intake of dairy cows was estimated using the n‐alkane technique. Cultivar differences for all sward structural characteristics were found except for bulk density and tiller density in 2003. Cultivars differed for proportions of pseudostem, stem (in 2003 only) and dead material. The chemical composition of the herbage was different among cultivars, with the water‐soluble carbohydrate concentration showing large variation (>0·35). Cultivars differed in susceptibility to crown rust. Herbage intake differed among cultivars in 2002 (>2 kg DM) but not in 2003. Herbage intake was positively associated with sward height, DM yield and green leaf mass. Canopy morphology did not affect herbage intake. Crown rust affected herbage intake negatively. It was concluded that options for breeders to select for higher intake were limited. High‐yielding cultivars and cultivars highly resistant to crown rust were positively related with a high herbage intake.     

In vitro fermentation patterns and methane production as influenced by cultivar and season of harvest of Lolium perenne L.

In some European countries, the majority of annual enteric methane (CH₄) emissions by ruminants occur at pasture – a direct result of the predominance of grazing within ruminant production systems. However, there are only limited data available as to the effect of perennial ryegrass cultivar and season of harvest on CH₄ production. Using the in vitro gas production technique, the effect of perennial ryegrass cultivar on fermentation characteristics and CH₄ production was determined (Experiment 1) and the persistence of these traits throughout the growing season for two cultivars, identified from Experiment 1 as having either a high or low methanogenic potential, was examined (Experiment 2). In Experiment 1, organic matter (OM) digestibility and cumulative total gas production profiles were unaffected by cultivar but, with regard to the kinetics of CH₄ production, the asymptote value (A), cumulative CH₄ yield at 72 h, and the fractional rate (μ) of CH₄ production at both time of 0·5A(T)(μCH₄T) and at 48 h (μCH₄48h) were significantly (P < 0·05) different. The amount of digested OM, as a proportion of cumulative CH₄ production (DigOM/CH₄) at 24 and 72 h after commencement of inoculation, revealed that the amount of substrate required to produce 1 ml of CH₄ also differed significantly between cultivars (P < 0·01). In Experiment 2, regrowth number significantly modified the majority of measured samples (P < 0·01); cultivar effects were limited to the lag phase of the cumulative CH₄ production curve and DigOM/CH₄ at 8 h only (P < 0·05). These results suggest that differences exist between cultivars in how OM is partitioned following microbial fermentation and that these differences demonstrate persistency throughout the growing season. In the course of time it may be possible to exploit these differences through cultivar selection and plant breeding programmes, and thereby reduce enteric CH₄ emissions within pastoral production systems.     

Between-cultivar variability in breaking force of components of perennial ryegrass (Lolium perenne)

The variation in force required to break the leaves and pseudostems of perennial ryegrass ( Lolium perenne ) was measured on twelve New Zealand commercial and pre-release cultivars using a punch and die apparatus which enabled up to 100 simultaneous breaks per sample. Perennial ryegrass cultivars were sampled from 2-year-old pure swards in two studies carried out in spring and autumn. In the first study, the forces required to break old and young leaves, and pseudostems, of six cultivars ranging in heading date and ploidy were compared at a designated morphological stage during spring growth. Force was measured as Newtons (N) mg⁻¹ dry matter (DM) and the difference between the force required to break leaves and pseudostems was significant ( P  < 0·01). The youngest emerged leaf required a greater force by 0·13 than the oldest green leaf (5·60 and 4·90 N mg⁻¹ DM for young and old leaf respectively). The force required to break leaves (average of young and old leaves) of 5·25 N mg⁻¹ DM was 0·12 greater than the force required to break pseudostems (4·59 N mg⁻¹ DM). This difference was consistent across most cultivars. In the second study, the force required to break leaves of ten cultivars was tested daily over three autumn periods when all cultivars were sampled at a regrowth age of 2 weeks. The cultivar Nevis was used in each period whilst the other cultivars were used in one period. When averaged across periods, cultivars could be separated into three groups of high, medium and low forces required to break leaves. The difference between the means of the high and low groups was 0·33 with a range from 3·29 to 4·91 N mg⁻¹ DM.     

Effect of temperature regime on germination of seed of perennial ryegrass (Lolium perenne)

Perennial ryegrass ( Lolium perenne L.) is one of the most valuable temperate pasture grasses in the world. Understanding seed germination in response to temperature can help determine its optimal seeding times. Germination tests for two cultivars of perennial ryegrass (cv. 'Turfstar' and cv. 'Transtar') were conducted by applying thirty-six constant and alternating temperature regimes, ranging from 5 to 40°C (16 h night/8 h day). Responses in germination rate to these temperature regimes were then used to construct a quadratic response surface, giving estimated germination rates with confidence intervals at P  ≤   0·05 for both cultivars. Optimum germination rate occurred over a range of regimes: 15/25, 20/25, 20/30 and 25/30°C. Furthermore, a wide range in diurnal alternating temperatures resulted in maximum seed germination rate for cv. 'Transtar', while no constant temperature regime supported maximum germination rate for cv. 'Turfstar'. A temperature-dependent germination rate function for perennial ryegrass was developed which could be used in the future to predict location-specific optimum seeding times when national or regional monthly temperature data are available.     

Differences in winter-hardening between phenotypes of Lolium perenne with contrasting water-soluble carbohydrate concentrations

Carbohydrates are essential for the winter-hardening of temperate grasses, affecting both rate of hardening and level of frost hardiness. The rate and extent of winter-hardening of contrasting phenotypes of two Lolium perenne cultivars targeted for differences in water-soluble carbohydrates concentration were quantified under controlled hardening conditions and standardized freeze testing. The phenotype with the lowest water-soluble carbohydrates concentration had a lower rate of hardening and tolerated less frost than the other phenotypes but the effect of water-soluble carbohydrate concentrations on the hardening processes levelled off at increasing water-soluble carbohydrate concentrations, indicating a threshold. Contrasting responses to freezing were found for survival and for potential for tillering and regrowth. Mortality was little affected by the higher test temperatures, while tillering was negatively affected, even at −5°C. Compensatory regrowth of tillers surviving the lower test temperatures partially compensated for the effects of loss of tillers on regrowth. The hardiest phenotypes had a rapid lowering of LT₅₀ during the first 3 d (−1·65°C d⁻¹), acquiring LT₅₀ values at −15°C or below in 10 d. The hypothesis that rate and maximum level of hardening increase with water-soluble carbohydrates concentrations in stubble is supported by the results.     

Genotypic variation in patterns of root distribution, nitrate interception and response to moisture stress of a perennial ryegrass (Lolium perenne L.) mapping population

Genotypic variation in patterns of root distribution, nitrate interception and response to moisture stress were assessed in both parents and 198 progeny of a perennial ryegrass (Lolium perenne L.) full‐sibling mapping population. This was carried out in metre‐deep tubes of sand culture in a glasshouse experiment. The proportion of root dry matter (DM) weight in the top 10 cm of sand ranged from 0·33 to 0·75 and values of log₁₀(1 ? K), where K is the constant for an exponential model relating root DM weight and root depth, also showed wide variation among genotypes. The proportion of a pulse of ¹⁵N recovered in whole plants ranged from 0·124 to 0·431. There was a positive linear correlation between the proportion of ¹⁵N recovered and plant total DM weight, but no relationship between nitrate interception and patterns of distribution of DM weight of roots. Some genotypes responded to moisture stress by increasing root growth, and in others root growth was inhibited. It is concluded that this below‐ground variability in root variables may be an evolutionary adaptation by plant populations to survive heterogeneity in soil biotic and edaphic factors.     

Effects of Neotyphodium endophytes on growth, reproduction and drought-stress tolerance of three Lolium perenne L. genotypes

Three Lolium perenne L. genotypes collected from different natural habitats were tested for the effects of their fungal endophyte Neotyphodium spp. on plant growth and seed yield. Half the clones of the originally infected plants were subjected to fungicide treatment to eradicate the endophytes. In an experiment, the clones were planted separately into pots and were either watered adequately or subjected to drought stress. In the genotype collected from a dry site, the endophyte infection reduced plant growth at an adequate water supply, but increased regrowth under drought. In the genotype from a periodically either flooded or dry site, endophyte infection significantly promoted the development of reproductive tillers and seed production (effects which are associated with adaptation to drought). In contrast, the genotype that originated from a wet site showed higher sensitivity to drought stress when endophyte infection was present. The results suggest that environmental conditions in the original habitat of the plants may influence the symbiotic interaction between plant and fungus, probably through natural selection. However, endophyte‐induced increases in root dry weight and root/shoot ratio were recorded for all three genotypes. These features could be beneficial for plant persistence, especially on sites where water is the growth‐limiting factor.     

Longevity of seeds of Poa pratensis and Lolium perenne as affected by simulated soil tillage practices and its implications for contamination of herbage seed crops

In grass-seed production, the purity of the product is of major importance and high purity can only be obtained by preventing contamination with other species and cultivars during the whole crop rotation. In this context, seeds of previously grown grass species or varieties are particularly important. This study investigated the influence of different soil tillage practices on persistence of newly shed seeds of two grass species, Poa pratensis and Lolium perenne . Leaving the seeds at or close to the soil surface reduced seed survival of both species to low levels compared with strategies that involved a deep incorporation of the seeds shortly after seed-shedding. Survival rate of seeds of P. pratensis after 1 year was reduced to a low level when seeds were left at the soil surface for 2–3 months before incorporation. Seeds of L. perenne were less persistent and survival rate after 1 year was generally very low if the seeds were left near the soil surface for approximately 1 month before deeper incorporation. Use of the stale seedbed technique to reduce the seedbank of the two species in the autumn ahead of a spring-established grass-seed crop was also found to be effective in reducing the potential risk of contamination.     

Identifying causes of low persistence of perennial ryegrass (Lolium perenne) dairy pasture using the Basic Grassland model (BASGRA)

Recent years have seen a decline in herbage production and tiller populations in New Zealand's perennial ryegrass (Lolium perenne) dairy pastures. One hypothesis is that modern genotypes are less suited to the warmer, drier weather experienced under changing climate patterns. In this study, a combination of long-term trial data (2011–2017) and a process-based pasture model (BASGRA) was used to explore the causes and possible mitigation of the observed production and population loss at three sites (dryland sites in Northland and Waikato and an irrigated site in Canterbury). Bayesian calibration was used to identify the model parameter sets that were consistent with the trial data and to identify differences in plant morphology and responses between sites. The model successfully simulated the observed differences in tiller numbers between the dryland sites, where populations and production declined rapidly after the second year and the irrigated site where populations and production were maintained. Analysis of the model calibrations along with preliminary scenario simulations suggests that increased tiller mortality associated with drought was the main cause of persistence failure at the dryland sites and that decreasing grazing pressure or breeding for tolerance to higher temperatures may not be successful in preventing this.     

A new descriptive scale for early reproductive development in Lolium perenne L.

Early reproductive development in Lolium perenne L. has been widely used for defining growth stages in spring, and this has been useful in managing seed crops and in trials involving the use of growth retardants.
A numerical scale for apical development based on morphological criteria, has been successfully used in research. The scale identifies eleven clearly recognizable stages of apical development, from the vegetative apex through to the immature, but completely-formed spike. It is intended that the key should be used to identify consistantly precise stages in apical development.     

Concentration of ergot alkaloids in Czech ecotypes of Lolium perenne and Festuca pratensis

The content of ergot alkaloids (ergovaline and chanoclavine), and their production in October 1996 and during the period May–September 1997, were investigated in seventeen ecotypes of perennial ryegrass ( Lolium perenne L.) and in nineteen ecotypes of meadow fescue ( Festuca pratensis Huds.), naturally infected with Neotyphodium spp. The ecotypes were collected in the north-eastern part of the Czech Republic. In 1996 the content of ergovaline in the ecotypes of perennial ryegrass ranged from 0·00 to 2·73 μg g^–1 dry matter (DM) (one cut), and in 1997 from 0·00 to 4·65 μg g^–1 DM (five cuts). In meadow fescue the content of ergovaline varied from 0·00 to 0·61 μg g^–1 DM (one cut) in 1996, and in 1997 from 0·00 to 2·31 μg g^–1 DM (five cuts). The content of chanoclavine (investigated in 1997 in four cuts only) in perennial ryegrass ranged between 0·00 and 3·39 μg g^–1 DM, and in meadow fescue between 0·00 and 2·26 μg g^–1 DM. Most ecotypes of L. perenne reacted to the high temperature and heavy rainfall in June and July of 1997 with an enhanced production of ergovaline, whereas the content of chanoclavine was not changed. Such reaction to stress conditions was not observed in the ecotypes of F. pratensis. Large differences in the production of both ergot alkaloids between different ecotypes of both plant species were observed.     

黑麦草生产能力及其抗热性评价

综述了黑麦草农艺性状及营养成分两方面的生产能力及差异,总结了黑麦草的育种方法,指出了高温胁迫对黑麦草的危害以及黑麦草抗热性的鉴定方法,提出了今后黑麦草抗热性育种的研究方向。     

Fatty acid composition and nutritive value of twelve cultivars of perennial ryegrass

A comparison was made of the fatty acid composition and nutritive value of twelve cultivars of perennial ryegrass ( Lolium perenne L.) differing in heading date and ploidy level. The cultivars were sown in triplicate plots and three sequential cuts of herbage were taken at 20-d intervals during the late spring and early summer to describe the fatty acid composition and other measurements of nutritive value. Differences between cultivars were recorded for DM content ( P  <   0·01) and concentrations of gross energy, neutral-detergent fibre (NDF) and ash ( P  <   0·05). Diploid cultivars had a higher DM content and concentration of NDF ( P  <   0·01) than tetraploid cultivars whilst late-heading date cultivars had the lower NDF ( P  <   0·05), ADF and ash ( P  <   0·01) concentrations. There was variation between cultivars in fatty acid composition. Diploid cultivars had a higher concentration of C18:0 ( P  <   0·01) and C18:1 ( P  <   0·05) than tetraploid cultivars and late-heading date cultivars had the highest concentrations of total fatty acids, C16:0, C18:2 and C18:3 ( P  <   0·05). This was predominantly due to the cultivar Tyrella which is a diploid, late-heading cultivar. The study showed that some variation exists between perennial ryegrass cultivars in concentrations of polyunsaturated fatty acids, which may present opportunities to select for this trait, but further research on the developmental stages and degree of leafiness of cultivars is first required.     

Effects of an established sward of Lolium perenne L. on the growth and development of Rumex obtusifolius L. seedlings

Seedlings of Rumex obtusifolius L. were grown in gaps (11 cm diameter) in an established sward at Lolium perenne L., Root competition from the surrounding sward was controlled by PVC tubes and shoot competition was controlled by cutting the sward frequently. All combinations, with and without root competition and with and without shoot competition, were used.
Plants received either 40 or 80 kg N ha⁻¹ month⁻¹ and 0 or 300 kg K₂O ha⁻¹ and were harvested 39 d after emergence; a second harvest was made 12 d later, after shading to 20% of full sunlight.
The shoot dry weight per plant of R. obtusifolius was much more affected by root competition than by shoot competition, especially at low nitrogen applications. Potassium supply had no effect.
More dry matter was allocated to the leaf petioles under shaded conditions and the specific leaf area was greater. This morphological plasticity enabled R. obtusifolius to grow well under shading and to be affected only slightly by shoot competition from L. perenne.
The results suggest that competition for nitrogen was the main factor limiting the initial grown of R. obtusifolius in a ryegrass sward and that the morphological characteristics of R. obtusifolius make it less susceptible to competition for light in the early stages of its development.     

Morphology and nutritive value of perennial ryegrass cultivars at different phenological stages

Eight perennial ryegrass cultivars (representing the variations in ploidy, heading date and water‐soluble carbohydrates concentration) were investigated for morphology and nutritive value at three phenological stages: pre‐heading vegetative, reproductive and post‐flowering vegetative stages. Chemical compositions and digestibility of morphological components (lamina, pseudostem and reproductive stem) from each perennial ryegrass cultivar were analysed in a split‐plot design. At the vegetative stages, perennial ryegrass cultivars differed significantly in the proportions of lamina and pseudostem. Tetraploid cultivars, Base and Bealey, always had the greatest lamina proportion (51.8% and 53.2% at the pre‐heading and post‐flowering vegetative stages respectively). At the reproductive stage, the emergence of seed heads diminished the differences in morphology among cultivars. Perennial ryegrass cultivars also had distinct nutritive value throughout three phenological stages. The high‐sugar cultivar, AberMagic, had high WSC concentrations (276, 227 and 90 g/kg DM at the pre‐heading vegetative, reproductive and post‐flowering vegetative stages respectively); the intermediate‐heading cultivars, Kamo and Commando, generally had a lower organic matter digestibility in dry matter than the late‐heading cultivars at the pre‐heading vegetative (70.7% vs. 74.4%) and reproductive stage (63.9% vs. 68.2%). However, although the morphological components were different in nutritive value consistently, the differences in morphology did not account for the variation in nutritive value among perennial ryegrass cultivars.     

香根草、黑麦草和高丹草光合生理生态日变化的比较研究

采用LI-6400便携式光合仪测定了香根草、黑麦草和高丹草的叶片光合日变化以及某些生态指标。结果表明：3种植物的净光合速率Pn日变化呈双峰曲线,但峰值出现的时间不同,并且有野午休冶现象。香根草、黑麦草和高丹草的日平均Pn值分别是11.513、16.737、20.565μmolCO2·m-2·s-1曰香根草和黑麦草的蒸腾速率Tr呈双峰曲线,高丹草呈单峰值曲线;3种植物的Pn、Tr、Vpdl及高丹草的气体导度Gs与空气相对湿度RH呈显著负相关(-0.826＜r＜-0.994)、与Tl和PAR则呈显著正相关(0.714＜r＜0.995)。     

The influence of ethofumesate on the growth and development, chlorophyll and nitrogen content of Lolium perenne L.

Pot experiments and a hand-weeded field trial were carried out to investigate the direct effect of ethofumesate on the growth and development of Lolium perenne cv. S24.
In the field trial, ethofumesate was applied in autumn as Nortron 20% ec formulation in a volume of 2001 ha^-1 at rates of 0, 1 or 2 kg a.i. ha^-1. Treatments had no direct effect on yield in the following summer. In both the field trial and pot experiments there was initial growth suppression in the form of a reduction in shoot dry weight, lamina dry weight, tiller number per plant and plant height. However, ethofumesate increased the chlorophyll and nitrogen contents of L. perenne laminae, but only when available soil nitrogen was not limited.
It is concluded that ethofumesate can increase the yield of L. perenne by suppressing weed competition, but there is no evidence of the herbicide directly increasing the growth or yield of cv. S24.     

Seed development and seed shedding in North Italian ecotypes of Lolium multiflorum

Seed development and seed shedding were studied in seven populations of Lolium multiflorum. Five of these populations were from the North Italian Plains, two from the Piedmont area and three from the Lombardy region. The remaining populations were Delecta, a variety derived from ecotypes from the Lombardy Plain, and RvP Lemtal. Seed shedding was greatest in the three ecotypes from the Lombardy Plain and lowest in RvP Lemtal. The two ecotypes from the Piedmont Plain, and Delecta, gave intermediate values. Moisture loss during ripening was greatest in the Lombardy ecotypes but there were no differences in the time taken to reach maximum 1000-seed weight. Germination values had reached a maximum 27 d after anthesis in all populations. It is suggested that the North Italian ecotypes, or varieties derived from them, should be harvested by direct combining at a moisture concentration of 450 g kg^-1 (≡45%) to obtain the maximum weight of germinable seed.     

蓖麻花序特征及花芽分化的初步研究

本文对蓖麻花序特征、顶芽分化及内源IAA和ABA含量变化进行初步研究,结果表明：不同品种（系）间雌花率及雌花密度均存在极显著差异。蓖麻雌花并非完全单性花,各品种（系）雌花均存在退化雄蕊痕迹,但其发生率不同,单雌后代品系明显高于品种材料。单雌花序与两性花序的雌花中均存在退化雄蕊,且无显著差异,但花序中下部与上部之间差异显著。顶芽石蜡切片观察表明：蓖麻在5叶期以前,其顶芽基本处于叶芽生长阶段;在6-9叶期,顶芽逐渐进入花芽分化、生长阶段,而此阶段,顶芽内源IAA、ABA含量变化也十分明显。不同性别花序和花的内源激素含量测定结果显示：IAA含量,单雌花序＞雌花＞两性花序＞雄花;ABA含量,雄花显著高于雌花,但两性花序却明显低于单雌花序;IAA/ABA值,雌花＞单雌花序＞两性花序＞雄花;说明IAA含量和IAA/ABA相对含量可能在花性分化中具有较为重要的作用。