The influence of light and temperature during winter on growth and death in simulated swards of Lolium perenne

Two growth experiments were carried out in January-March 1978 in which simulated swards of perennial ryegrass were (1) heated by soil warming cables to give soil surface temperatures of approx. 7·5-9·C above ambient temperature, (2) shaded with netting to reduce light levels by approx. 50%, or (3) both heated and shaded.
Heating alone increased leaf appearance, death, extension, lamina size, leaf area index (LAI), tillering (month 1)and whole plant weight (month 2) and reduced stubble water-soluble carbohydrates and specific leaf weight (SLW).
Shading alone increased leaf extension, lamina size and LAI but to a lesser extent than did heating. Shading decreased SLW, leaf death rate, tillering (month 2), stubble carbohydrates and whole plant weight, but not herbage weight.
The effects of heating plus shading were similar to those of heating alone, except that the increases in leaf size, extension and LAI were even greater, and shoot bases and roots had low or negative growth rates.
In general the heating treatments caused a rapid turnover of leaf material, but net herbage growth was relatively insensitive. It is concluded that (1) temperature rather than light was limiting whole plant growth, especially from mid-February to mid-March and (2) mild, dull weather in winter is likely to induce tiller death associated with reduced investment in carbohydrate reserves.     

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.     

Interaction between water-soluble carbohydrate reserves and defoliation severity on the regrowth of perennial ryegrass (Lolium perenne L.)-dominant swards

A field-study was undertaken in Hamilton, New Zealand to determine if there was an interaction between water-soluble carbohydrate (WSC) reserve content and defoliation severity on the regrowth of perennial ryegrass-dominant swards during winter. Perennial ryegrass plants with either low or high WSC content were obtained by varying the defoliation frequency. At the third defoliation at the one-leaf stage and at the first defoliation at the three-leaf stage (harvest H₁), swards were mown with a rotary lawnmower to residual stubble heights of 20, 40 or 60 mm. All swards were then allowed to regrow to the three-leaf stage before again defoliating to their treatment residual stubble heights (H₂). Frequently defoliated plants contained proportionately between 0·37 and 0·48 less WSC in the stubble after defoliation, depending on the severity of defoliation. There was no interaction between WSC content and defoliation severity for herbage regrowth between harvests H₁ and H₂. Herbage regrowth was lower from swards containing low WSC plants compared with high WSC plants (2279 vs. 2007 kg DM ha⁻¹). Furthermore, swards defoliated to 20 or 40 mm had greater herbage regrowth compared with those defoliated to 60 mm (2266, 2249 and 1914 kg DM ha⁻¹ for swards defoliated to residual stubble heights of 20, 40 and 60 mm, respectively). Regrowth of perennial ryegrass was positively correlated with post-defoliation stubble WSC content within defoliation severity treatment, implying that WSC contributed to the defoliation frequency-derived difference in herbage yield. However, the effect of defoliation severity on herbage regrowth was not associated with post-defoliation stubble WSC content.     

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。     

Seasonal variation in growth characteristics of Lolium perenne and Trifolium repens in swards under different managements

In a range of perennial ryegrass/white clover swards, variation in the surface heights of the grass and clover components, the rates of increase of these surface heights and the specific leaf areas (SLAs) of ryegrass and white clover were described for 1 year. The swards were of an early- or late-flowering (Aurora or Melle respectively) perennial ryegrass variety growing with either a small- or a medium-leaved (Kent or Milkanova) white clover and were either continuously stocked by sheep or continuously stocked apart from a rest period in April-May (Aurora) or May-June (Melle).
The surface heights of grass and clover were not affected by the variety of their companion species, and the surface heights of the two clover varieties were similar. The grass was always taller than the clover, although the magnitude of the difference between the species varied with time of year and the timing of the rest period.
Before the summer solstice the rate of increase in height of grass was greater than that of clover except at cool temperatures (5°C) and warm temperatures (16°C), and in the unrested Melle sward. After the solstice the rates of increase in height, particularly of clover, were lower than the rates seen at similar temperatures before the solstice.
Overall, the SLAs of both clover varieties were greater than those of ryegrass when grown with Aurora but not when grown with Melle, and the SLAs of both species increased during the year. By October the SLAs of both grass varieties were less than those of their companion clovers.
The results are discussed in relation to their implications for the species composition of the swards.     

The influence of crown rust (Puccinia coronata) on tiller production and survival of perennial ryegrass (Lolium perenne) plants in simulated swards

Populations of Aberystwyth S23 and S24 ryegrass tillers were individually labelled and harvested at intervals to follow their life histories and productivity in response to the effect of infection with crown rust at two levels of nitrogen fertilization.
The number of tillers in a population was reduced as a result of infection with crown rust. Increased rates of tiller death due to the pathogen occurred throughout the population age-range and were most pronounced in the youngest tillers. Population changes were evident following infection and continued to be apparent in the following spring. The disease caused alterations in sward age structure, leading both to a higher proportion of older tillers in the more rust-damaged treatments and to a decrease in the weight of the herbage produced. It is concluded that rust infection of a sward may lead to changes in population structure with long-term yield effects, in addition to the immediate direct yield reductions commonly reported. The significance of these results to the ecological and agronomic balance of grassland is discussed.     

Canopy height measurements and non‐destructive biomass estimation of Lolium perenne swards using UAV imagery

In perennial ryegrass breeding programmes, dry‐matter yield (DMY) of individual plots is monitored destructively at the different cuts or derived from non‐destructive canopy height measurements using devices like rising plate meters (RPM). These approaches both have constraints. Destructive sampling implies low temporal resolution, restraining the study of dry‐matter accumulation rates, while RPM measurements are influenced by the canopy structure and limit intra‐field variability identification. We present a phenotyping methodology, based on the use of an affordable RGB camera mounted on an unmanned aerial vehicle (UAV), to monitor the spatial and temporal evolution of canopy height and to estimate DMY. Weekly flights were carried out from April to October above a field comprising a diverse set of accessions. To test the capacity of UAV imagery to estimate canopy height, 8 ground control points and 28 artificial height references were placed at different locations. Accurate flights with an RMSE as low as 0.94 cm were achieved. In addition, canopy height was recorded using an RPM and destructive biomass samples were collected. Different models (linear, multiple linear, principal components, partial least squares regression and random forest) were used to predict DMY, and their performance was evaluated. The best estimations were obtained by combining variables including canopy height, vegetation indices and environmental data in a multiple linear regression (R² = .81). All models built using UAV data obtained a lower RMSE than the one using RPM data. The approach presented is a possibility for breeders to incorporate new information in their selection process.     

Effect of Lolium perenne sward density on productivity under simulated and actual cattle grazing

The objective of this study was to quantify the effect of perennial ryegrass (Lolium perenne L.; PRG) sward density on seasonal and total DM yield under simulated grazing and animal grazing by cattle, and to assess the effectiveness of visually estimated ground scores (GSs) for predicting sward PRG density. The study incorporated five different seeding rates of PRG, each replicated three times, to simulate swards ranging in PRG density typical of different ages and conditions. There was no significant difference between defoliation managements for total DM yield, but sward PRG density had a significant effect on both the seasonal and total DM herbage yield under both systems. Under simulated grazing, total DM yield ranged from 10·7 to 12·0 t DM ha^?1 with increasing sward PRG density at a GS range of 1·70–4·28 (mean of 2 years’ data, P < 0·01). Under animal grazing, the yield range was from 10·3 to 12·2 t DM ha^?1 for a GS range of 1·50–3·39 (mean of 2 years’ data, P < 0·01). The largest differences in DM yield occurred during the spring period. The relationship between sward DM yield and GS was significant (P < 0·001) for both simulated and animal‐grazed swards. Each unit increase in midseason GS (June) related to an average yield increase of 350 kg DM ha^?1 under simulated grazing and a 721 kg DM ha^?1 increase under cattle grazing. Every unit increase in the GS at the end of the grazing season (December) was associated with a 460 or 1194 kg DM ha^?1 increase under simulated and animal grazing, respectively. These results show that visual estimates of density were an effective tool in describing PRG density and that this could be related to DM yield potential. Further investigations may provide a threshold value below which the renewal of swards could be advised based on a visual GS of PRG.     

Effects of vernalization on the subsequent rates of leaf extension and photosynthesis of perennial ryegrass (Lolium perenne L.)

Temperature and daylength treatments have been used to investigate the apparent link between the potential for rapid leaf extension in spring and floral initiation and development in Lolium perenne (cv. Vigor). After growth at 5°C for 10 weeks (vernalization treatment) the leaves which developed at 15°C expanded significantly faster than non-vernalized controls under both long and short days. Under long days floral development was initiated in vernalized plants but under short days it was not. Specific leaf area was higher in vernalized than non-vernalized plants under long days.
The photosynthetic capacity of leaves which developed on plants which had been previously vernalized and then grown at 15°C under long days was higher than non-vernalized controls when expressed on a leaf area and dry weight basis, but not on the basis of chlorophyll content. Vernalization followed by short days resulted in a smaller and non-significant increase in the photosynthetic capacity of leaves.
Application of gibberellic acid (GA) as a spray to non-vernalized plants did not affect floral development but did increase the rate of leaf extension. The application of GA did not affect specific leaf area or photosynthetic capacity expressed on an area basis.     

Variation in inflorescence characters and inflorescence development in ecotypes and cultivars of Lolium perenne L.

A sufficient database of seed yield components is the prerequisite to predict which component contributes most to the seed yield complex and to dissect the biology of the complex quantitative trait seed yield. Phenotypic variation of eleven vegetative and reproductive traits was characterized for 2481 individuals from fifty Lolium perenne L. (perennial ryegrass) ecotypes and cultivars. Considerable levels of among‐ and within‐population variation were found across several inflorescence characters. Principal component and canonical variate analysis separated ecotypes from cultivars, and cultivars generally had later dates of ear emergence, better spring and summer growth, longer rachis length and more spikelets per spike than ecotypes. Strong positive relationships were seen between inflorescence characters using both correlation and regression analyses. The strong relationship between rachis length and other inflorescence characters suggested that rachis length could be used as a performance predictor for several other characters of the spike. This phenotypic study has helped to determine basic patterns of morphological diversity and correlations between characters. It is discussed in which way further developmental genetic studies can be directed.     

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.     

The use of growth regulator RSW0411 in seed crops of Lolium perenne L.

Field trials were conducted in 1985 and 1986 to assess the effects of the growth regulator RSW0411 on a Lolium perenne L. (perennial ryegrass) seed crop. Applications were made at spikelet and floret initiation, and at 1, 2, 3 and 4 kg active ingredient ha⁻¹.
Seed yield increases in 1985 were associated with increased reproductive tiller numbers and seed numbers, but in 1986 there were no seed yield responses.
Lodging was less severe in 1986, and there was less secondary filtering during seed maturation. In 1985, fertile tiller numbers were lower in control plots at harvest. These results are discussed in relation to the seasonal disparity in seed yield response.
RSW0411 application significantly decreased culm lengths by shortening the basal internodes, and this effect was greater with higher application rates. Stem width was increased in 1985 at the base and middle of the stem, and earlier applications increased the basal width by more than later applications.
Germination was not affected by RSW0411 treatment, except when applied at 4 kg active ingredient ha⁻¹ in 1985, when there was a small decrease. This was related to a low 1000-seed weight, and is discussed in relation to seed maturity at harvest.     

Nitrogen studies in Lolium perenne grown for seed III. The effect of nitrogen and water stress

The effects of water and nitrogen (N) availabilities and their possible interactions on the seed yield and yield components of S23 perennial ryegrass grown in pots in the glasshouse were investigated. Increasing the N fertilizer from 0 to 5 g per pot more than doubled the seed yield but no further increase in seed yield was found by increasing N fertilizer from 5 to 10 g per pot. The main effect of increasing water availability was to increase both seed and straw yield. When N had been applied, accumulated evapotranspiration was linearly related to seed and straw yield. The component of yield having the greatest effect on yield was the number of fertile tillers and, although both N and water availability affected this component, N had the larger effect. Implications of these responses are discussed in relation to the potential yield of the crop and to the field situation.     

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.     

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.     

Factors influencing the proportion of natural hybrids between Lolium perenne L. and Festuca pratensis Huds. in permanent pastures

Hybrids between ryegrass and meadow fescue form rarely but occur widely in old pastures. In one experiment six different hybrid genotypes were able to compete, on equal terms, with members of their parental species. In two others the proportion of the total yield due to one hybrid genotype was increased by added fertilizer or by reduced water supply.     

Sward height, structure and leaf extension rate of Lolium perenne pastures when grazed by overwintering barnacle geese

Abstract The grazing of agricultural pastures during winter and spring by geese is considered an important agricultural problem in parts of the U.K. This study describes the sward structure, leaf extension and senescence rates of Lolium perenne‐dominated pastures that are frequently grazed by barnacle geese (Branta leucopsis) during winter in South‐west Scotland, as well as the conducting of a field experiment that simulated grazing to investigate the effects of defoliation. Gross leaf extension and senescence rates were strongly related to temperature, daylength and average tiller size, resulting in positive values of net leaf extension rate per tiller for most of the winter. Total tiller length declined from November to January but increased from January to April. Sward height, however, declined consistently from October to April, suggesting that swards were becoming trampled by repeated visits by flocks of geese over this time. The structure of individual tillers was found to vary slightly over the winter, with tillers becoming more dominated by younger leaves towards the end of the winter. Experimental defoliation of tillers suggested that absolute leaf extension rates did not respond in an under‐ or over‐compensatory manner, even when tillers were nearly completely defoliated. The results suggested that sward structure and leaf extension rates are not unduly affected by repeated grazing by overwintering geese and that short‐term depletion and trampling are the main impacts.     

The effect of light intensity after anthesis on dry matter distribution and seed yield of Lolium perenne L.

The effect of light intensity after onset of anthesis on dry matter distribution, water soluble carbohydrate (WSC) concentration and seed filling in Lolium perenne was investigated in a pot experiment. Spaced plants of five clones were brought to flowering and exposed to four light intensities (24%, 57%, 100%, 115% of natural light) from 7 days after onset of anthesis onwards. The two oldest flowering tillers (i.e. the main and the first tiller) were separated from the rest of the plant and dissected after a seed-filling period of 450°Cd (temperature sum counting from 7 days after onset of anthesis). Their stem dry weight and WSC concentration were reduced at the lowest light intensity. WSC amount was more reduced in the first, younger, tiller. Seed yield per ear of the main tiller was reduced by 14% because seed set and average seed weight were lower at the 24% light intensity. Seed yield and seed number per ear of the first tiller were lowered by 21% and 17%, respectively, at the 24% light intensity, but average seed weight was not. The relative contribution of the seeds to total tiller weight was about 20% and was similar for both tillers and all light intensities. WSC concentration and WSC amount in the stem + rachis were reduced only at 24% light in the main tiller and at 24% and 57% light in the first tiller. It is inferred that seed yield was not limited by assimilate availability, but by the ability of the seeds to utilize fully the abundantly available reserves in the stem. The tiller can support seed yield under a wide range of tight intensities.     

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.