SEED PRODUCTION STUDIES ON PERENNIAL RYEGRASS, TIMOTHY AND PRAIRIE GRASS

This paper considers the sequence of seed development by measurement and observation of ehanges in seed moisture content, weight, colour, endosperm consistency, germination capacity and yield in perennial ryegrass Grasslands Ruanui, timothy Grasslands Kahu and a New Zealand strain of prairie grass (Bromus unioloides). Seed moisture content was high (60–70%) after anthesis and declined at different rates, depending on the species and the weather. Maximum seed dry weight (maturity) was reached approximately 30 days after peak anthesis in all species and was influenced by N application and grazing. Colour changes in the seedhead and endosperm solidification proved unsatisfactory as measures of maturity. Endosperm solidification was reached 4, 11 and 12 days before maximum seed dry weight in ryegrass, timothy and prairie grass, respectively. Seeds removed from the plant as little as 7, 10 and 4 days after anthesis in ryegrass, timothy, and prairie grass, respectively, germinated, provided that dormancy was broken; but they did not retain viability after 3 months storage. Maximum viability of ryegrass, timothy and prairie grass seed tested immediately after harvest occurred 30, 38 and 16 days after anthesis, compared with 17, 35 and 12 days, respeetively, when germination testing was delayed for 3 months. In ryegrass, grazing caused a significant increase in seed yield in the second year while N application alone did not significantly increase yield. In timothy, grazing and to a lesser extent N, were both detrimental to seed yield in the second year. Highest yields were obtained in treatments not receiving either grazing or N. In prairie grass, grazing depressed and N application increased yields in both years. In all species similar or increased yield could be obtained by mowing the crop at, or slightly prior to, seed maturity and allowing seed development and ripening to continue on the cut straw for up to 10 days. Using this technique higher seed yields were obtained in ryegrass and timothy than when seed was direct-harvested. In prairie grass similar yields were obtained whether the erop was direct-harvested or threshed following drying in the swath for 10 days. The use of ‘drying curves’ for the prediction of correct cutting time of seed crops is suggested for ryegrass, timothy and prairie grass.     

Productivity of prairie grass (Bromus willdenowii Kunth) affected by sowing date and the head smut fungus (Ustilago bullata Berk.)

Effects of sowing date, fungicide seed treatments, and the head smut fungus Ustilago bullata Berk, on establishment, herbage, and seed production of prairie grass cv. Grasslands Matua ( Bromus willdenowii Kunth ) were measured in autumn- and spring-sown field trials in New Zealand.
Autumn seedling establishment from thiram-treated seed relative to untreated seed was increased by 27% from the earliest sowing (early March), and by 65% from the latest sowing 8 weeks later. Infestation of seed with ustilospores of U. bullata reduced seedling establishment at all but the latest autumn sowing. Fungicide seed treatments had little effect on spring seedling establishment, but V. bullata infestation of seed reduced establishment compared with healthy seed, particularly after late sowings (early November).
Herbage production was greater from early than from late sowings in both autumn and spring, and was less in plants grown from U. bullata infested seed than in plants from healthy seed. Enhanced autumn establishment from thiram-treated seed led to improved herbage production when plants were young, but this effect disappeared about 18 weeks after late autumn sowing. Overall herbage production from spring sowing was more than twice that from autumn sowing.
A 6-week delay in autumn sowing led to a 58% reduction in seed production the following summer.
Early autumn and spring sowings, together with adequate control of seedling diseases, are likely to give optimum herbage and seed production from prairie grass.     

THE EFFECT OF THE TIMING OF AMMONIUM NITRATE DRESSINGS ON THE HEADING OF S48 TIMOTHY (PHLEUM PRATENSE L.)

The response of single plants of Aberystwyth S48 timothy ( Phleum pratense L.) to increasing rates of ammonium nitrate applied in the spring and autumn is described.
All spring treatments markedly increased the number of heads, as well as the total head-length per plant, but autumn dressings had no significant effect. Percentage head-length distribution was also greatly influenced by applications of nitrogenous fertilizer in the spring. Each higher rate of application gave a proportionate increase in the number of heads over 9 cm. in length, and a corresponding reduction in the frequency of those of 4 cm. or less. There was a positive correlation between head-length and number of florets per unit length.
The results are discussed in terms of their relationship to current practices in managing timothy crops for seed.     

THE EFFECT OF DEFOLIATION ON THE SEED-PRODUCING CAPACITY OF BRED VARIETIES OF GRASSES

Leafy grass varieties S24 perennial ryegrass, S26 cocksfoot, S215 meadow fescue and S51 timothy were cut in October and at various growth stages in spring; S53 meadow fescue was grazed at similar periods. Sub-plots were given different applications of N. The control plots had similar manurial treatments but were not defoliated. The seed yield of S24 following October cutting was significantly lower than the control in the first harvest year and significantly higher in the second; the mean yields over 2 years, as a result of cutting at this stage and before ear formation in March, were very similar to those of the control treatment. The mean yields from 3 harvest years for S26, S215 and S51 were generally improved by cutting in October and before ear formation, but cutting after ear formation reduced the number of heads and the seed yields. Extra N applied in conjunction with the defoliation had no significant effect on the number of seed heads or on seed yield. Double grazing of S53 in Sept. and Dec. reduced significantly its seed production in both harvest years. Application of N in spring gave a greater response than a similar application in autumn.     

Persistence of tall fescue in a subtropical environment: tiller survival over summer in response to flowering control and nitrogen supply

Enhancing pasture persistence is crucial to achieve more sustainable grass‐based animal production systems. Although it is known that persistence of perennial ryegrass is based on a high turnover of tillers during late spring and summer, little is known about other forage species, particularly in subtropical climates. To address this question, this study evaluated survival of grazed tall fescue tillers growing in a subtropical climate. We hypothesized that hard tactical grazing during winter to remove reproductive stems (designated as ‘flowering control’), and nitrogen fertilization in spring, would both improve tiller survival over summer, and thus enhance tiller density. This was assessed in two experiments. In both experiments, few tillers appeared during late spring and summer and so tiller density depended on the dynamics of vegetative tillers present in the sward in spring. In Experiment 2, flowering control and nitrogen fertilization both enhanced the survival of that critical tiller cohort, but the effects were not additive. Responses were similar but not statistically significant in Experiment 1, which had a warmer, drier summer and lower overall survival rates. Unlike grasses in temperate environments, persistence of tall fescue in this subtropical site appeared to follow a ‘vegetative pathway’; i.e., new tillers were produced largely in autumn, from vegetative tillers that survived the summer.     

SEASONAL VARIATION IN APPEARANCE AND GROWTH OF GRASS ROOTS

Data from two field experiments, and from a root observation trench, suggest an annual cycle of root growth by grass swards. The production of adventitious roots from the tillers of four grass species, and from perennial ryegrass in a grass/clover sward varied greatly during the year. They were produced in increasing numbers through late winter to early spring; this rate fell sharply in April or May, and was low in summer. The differences in this annual cycle between grass species were small. Examination of roots of perennial ryegrass in a glass-sided trench showed that root elongation was most rapid in summer and was largely confined to the lower soil horizons. Rather more roots were produced in early autumn than during summer, but the rate of elongation was slow from October until spring.
Individual roots of grasses appeared to survive for limited periods. Their longevity depended upon the time of year in which they were first formed; those produced in autumn or winter lived longer than those produced in spring or summer.     

THE EFFECT OF COMPANION GRASS AND SEED RATE ON THE PRODUCTIVITY OF A TALL-FESCUE SWARD

The effects of sowing from 10 to 50 lb/acre of tall-fescue seed on plant establishment and on the annual and seasonal distribution of DM yield of tall fescue and its companion grasses were measured through two growing seasons. Increasing the seed rate increased plant numbers per unit area and decreased per cent establishment. Adding Scots timothy as a companion grass depressed both plant number and per cent establishment. S37 cocksfoot and New Zealand perennial ryegrass reduced both still further. Total annual dry-matter production was relatively unaffected by varying either the seed rate or the companion grass. The contribution of tall fescue to total production was highest when sown alone and was successively reduced by timothy, perennial ryegrass and cocksfoot. When sown alone there were only slight differences in contribution of tall fescue due to seed rate. With any companion grass the contribution from tall fescue increased with increasing seed rate. Early growth was not reduced by modifying seed rate but was reduced by all the companion grasses. It is concluded that increasing the seed rate of tall fescue above 30 lb is not justified, that Scots timothy can be sown at 3 lb/acre with tall fescue without affecting total production or early growth of the mixture, with the advantages of control of unsown species and improved palatability of the herbage.     

THE EFFECT OF CUTTING TIMOTHY (PHLEUM PRATENSE L.) GROWN FOR PRODUCTION OF SEED

An experiment is described in which timothy, grown for production of seed, was cut once annually on dates ranging from 23 October to 24 May. New tillers in selected plants were labelled at monthly intervals during autumn and winter for 2 years, to determine the pattern of production of tillers and to assess the reproductive development of categories of tillers. The majority of tillers were produced in autumn, but production was continuous throughout the winter. There was a big increase from the first to the second year in the total number of tillers produced, but neither the pattern nor amount of tiller production was affected by cutting. The % of fertile tillers was highest in the oldest tillers, and was affected little by the increase in the number of tillers from the first to the second year; cutting had little effect on the % fertility except where floral primordia were removed by the cut on 24 May. In all treatments at least 90% of the ears at harvest were produced by tillers initiated the previous autumn. The penetration of light to the base of the plants was increased considerably by cutting, but it is concluded that interception of light by the whole plant is more beneficial than an increase in the penetration of light into the plant and a concomitant reduction in the total interception of light by the plant.     

Primary growth and quality characteristics of Bromus willdenowii and Lolium multiflorum

Prairie grass ( Bromus willdenowii Kunth) and a tetraploid Westerwolds ryegrass ( Lolium multiflorum Lam.) were established in a field trial in April 1987 and grown for a 3-month period of undisturbed growth. During this period the biomass partitioning and forage quality of each plant component was compared for the two species. Leaf, tiller and plant populations were assessed on ten occasions while accumulation of herbage and roots, chemical composition, leaf area and light interception were determined on six occasions. Herbage was divided into leaf lamina, inflorescence, vegetative and reproductive pseudostem. Nitrogen, water-soluble carbohydrates, ash, cell wall and in vitro digestibilities were determined.
Prairie grass had lower plant, tiller and leaf populations but larger tillers and more live leaves per tiller than Westerwolds ryegrass. Both species had similar light interception and leaf area index. Roots were distributed more evenly and to greater soil depths in prairie grass. Leaf lamina made major contributions to herbage DM accumulation and accumulation of the various chemical components, but as reproductive development occurred, reproductive pseudostem became a major component of the total sward. Harvesting herbage to gain optimum quantities of DM, herbage quality and regrowth is discussed. It is concluded that prairie grass is a high-yielding, high-quality forage grass, comparable with Westerworlds ryegrass.     

Leaf dynamics of timothy and meadow fescue under Nordic conditions

Leaf production dynamics of timothy ( Phleum pratense L. cv. Tarmo) and meadow fescue ( Festuca pratensis Huds. cv. Kalevi) were studied in Finland under field conditions in mixed swards containing timothy, meadow fescue and red clover ( Trifolium pratense L. cv. Björn) harvested by cutting. Fifteen randomly selected tillers of both species were marked and leaf number, leaf appearance rate (LAR), leaf elongation rate (LER), leaf senescence rate (LSR) and leaf live span (LLS) were followed through weekly observations. Concurrently another set of fifteen tillers of each species was dissected on each occasion to determine the number of unemerged leaves and development stage of the apex. The first observation period was between 26 April and 22 June and the second between 22 July and 22 September.
The species differed clearly for most of the measured parameters but most differences were dependent on the season. In spring, timothy started growth earlier, had higher LAR (+57%), produced more leaves (+83%) with higher LER_gross (+58%), higher LSR (+61%), higher LER_net (+54%) and produced larger tillers (+193% in leaf area) than meadow fescue. In autumn, timothy still had a higher LAR (+80%), produced more leaves (+60%) with higher LER_gross (+36%), but had also a higher LSR (+77%) and thus similar LER_net (0%) to meadow fescue. Timothy tillers were only slightly larger than meadow fescue tillers (+20% in leaf area). The consequences of these findings on competition in mixed swards and implications for grazing theories are discussed.     

A HISTORY OF THE INTRODUCTION OF TIMOTHY AND COCKSFOOT INTO ALTERNATE HUSBANDRY IN BRITAIN

The first recorded sowing of American timothy seed in England was in 1743, but the event was otherwise without significance since nothing further was heard of this sowing. The stimulation of interest in timothy began with the sowing by Bartholomew Rocque in the late summer of 1763 of the seed received by the London Society of Arts from America. There was during the first years some confusion regarding its vegetative characters, and also in recognizing the difference between the flowering heads of timothy and those of meadow foxtail ( Alopecurus pratensis ). A hindrance to progress in the use of timothy arose, without doubt, from lack of experience, and lack of suitable implements to provide the fine, firm, clean seed-bed required. Midsummer sowing may have contributed to failure, for timothy establishes itself best in a moist soil. Timothy seems to have maintained a rather modest place during the 19th and early 20th centuries. The first real progress was made after 1920 when the agronomic potential of the valuable material among our indigenous populations of grasses was appreciated and developed into a number of new cultivars adapted to current needs.     

AN ASSESSMENT OF (2-CHLOROETHYL) TRIMETHYLAMMONIUM CHLORIDE AS A POTENTIAL AID TO GRASS-SEED PRODUCTION

The effects of (2–chloroethyl) trimethyIammonium chloride (CCC) on inflorescence production and culm length in seedlings of Aberystwyth S51 timothy ( Phleum pratense L.) are described. Concentrations of CCC ranging from molar to 0.01 molar were applied as a soil drench at a rate of 75 ml/sq. ft to plants at the 3rd, 6th or 8th leaf stage. All treatments reduced the culm length at heading, maximum effect being obtained from those given at the earliest growth stages. The number of heads per plant and the total head-length per plant increased with treatment, showing a maximum after molar treatments at the 6th or 8th leaf stages. Mean head-length remained relatively constant. The results are discussed in relation to seed production problems and the questions to be answered prior to field-scale treatment are outlined.     

THE PRODUCTION OF EARLY SPRING GRASS

The effects of autumn management and nitrogenous manuring on the production of early spring grass were studied from 1954 to 1957 on farms throughout England and Wales.
The most important management factor influencing spring yields was the application of spring nitrogen. In the early spring 6 to 10 lb. of dry matter were produced per lb. of nitrogen applied. Autumn nitrogen increased herbage yields in the auttimn by 8 to 17 lb. of dry matter per lb. of nitrogen applied.
Italian and perennial ryegrass responded well to autumn nitrogen but it was important to graze off the extra growth fairly quickly to prevent the sward becoming too open. The yield of ryegrass swards which received spring nitrogen were very much higher if they were grazed during the previous September and October than if they had been rested during those months.
In two years neither the autumn management nor alumni fertilizer appeared to influence the spring yields of cocksfoot, timothy or meadow fescue swards, in one year, however, when autumn growth was poor, autumn nitrogen and resting resulted in slightly better yields on these swards in the following spring.
It is suggested that resting swards during September and October to build up "reserves" is relatively unimportant in relation to the yields of grass in the following spring.     

THE INFLUENCE OF PLANTING DATE ON THE SEED-BEARING CAPACITY OF TILLERS OF PERENNIAL RYEGRASS SPACED PLANTS GROWN FOR SEED

A study was made of the fate of tillers arising at different times in the autumn and the contribution which these tillers made to the final seed yield of single spaced plants of perennial ryegrass planted at different dates for seed production. The flexibility in tiller function and in seed-bearing capacity exhibited within these plants resulted in October-planted material producing as much seed in the following harvest-year as August-planted material, although the latter had twice as many living tillers per plant at harvest. This reproductive versatility has economic importance and may have biological significance in certain ecological conditions.     

Herbage productivity of prairie grass, reed canary-grass and phalaris

The herbage production and quality of swards of three grass species, prairie grass (Bromus willdenowii Kunth), reed canary-grass (Phalaris arundinacea L.) and phalaris (Phalaris tuberosa L.) were compared with perennial ryegrass (Lolium perenne L.) and hybrid ryegrass (L. perenne L. ×L. muitiflorum Lam.) under 6–cut (experiment 1) and 4–cut (experiment 2) regimes over 3 years at Ayr; annually, 360 kg ha^-1 fertilizer N were applied. At Edinburgh prairie grass was compared with Italian ryegrass (L. multiflorum Lam.), timothy (Phleum pratense L.) and cocksfoot (Dactylis glomerata L.) under an annual 4–cut regime for 3 years (experiment 3); fertilizer N application totalled 350 kg ha^-1 annually. Prairie grass gave the highest annual dry matter (DM) production at Ayr, averaging 11·99 t ha^-1 in experiment 1 and 15·62 t ha^-1 in experiment 2. Reed canary-grass was much less productive whilst phalaris did not persist after harvest year 1. On average, prairie grass gave 8–10% more DM than the three ryegrasses in the 6-cut system but its advantage was much less under the 4-cut regime. In experiment 3, the DM production of prairie grass and Italian ryegrass were similar in year 1, but following winter damage prairie grass gave the lowest production in subsequent harvest years. Prairie grass had digestibility (OMD) values lower than the ryegrass but higher than reed canary-grass, timothy and cocksfoot. The water soluble carbohydrate (WSC) concentrations in prairie grass were markedly higher than in timothy and cocksfoot but lower than those in Italian ryegrass. Prairie grass had relatively low P and Mg concentrations. Reed canary-grass had relatively low OMD and Ca, but high N, P, K and Mg contents. It is concluded that prairie grass may have potential in the UK as a special-purpose species for conservation management but mainly in the milder climatic areas. The Phalaris species evaluated had disappointing agronomic potential.     

FERTILE TILLER PRODUCTION AND SEED YIELD IN MEADOW FESCUE (FESTUCA PRATENSIS L.)

The fertile tiller production and seed yields of spaced plants of meadow fescue varieties were found to vary widely with the time of sowing, particularly in the first harvest year. The benefit of early spring sowing is indicated in the total productivity over two harvest years.
Varieties differed in the amount of heading in the seeding year; the American type was the most prolific. This had a bearing on the relative yields in the first harvest year, since many of the potentially productive tillers of this variety were wasted through the plants becoming ripe to flower in the seeding year and, therefore, the rate of tiller production was checked. Competition within the plant appeared to be the cause of tillers remaining vegetative when conditions otherwise seemed suitable for generative development. The smaller-sized plants gave a higher proportional increase of fertile tillers at harvest than would be expected from the vegetative tillers counted in April, while at the same time there was a negative relationship at harvest from plants which had a very high number of vegetative tillers.
Infertility as reflected in a poor seed set at the time of harvest is a serious wastage of potential production in meadow fescue and, in selecting for superior seed output, breeders need to take this factor into consideration.
The 1000-seed weight varied between varieties, the pasture type being inferior to the two hay types.     

Productivity and persistence of prairie grass (Bromus willdenowii Kunth)

A field trial measured effects of the head smut fungus Ustilago bullata Berk, on forage productivity of prairie grass ( Bromus willdenowii Kunth). Simulated swards containing different proportions of U. bullata infected and non infected plants were established in the autumn, and sward and plant parameters were measured over the following 15 months. Total herbage produced from swards containing only non-infected plants was 27.3 t ha⁻¹, while that from totally infected swards was 14.6 t ha⁻¹. Infected plants produced fewer and lighter tillers than non-infected plants when both were growing together in swards. Almost all the U. bullata infected plants died during an epidemic of bacterial wilt disease (caused by Xanthomonas campestris pv. graminis (Egli, Goto and Schmidt) Dye), while most of the'non-infected plants survived. The deleterious effects of U. bullata on individual plant productivity affected sward productivity only when the proportion of infected plants in swards was greater than 50%. Plants not infected with U. bullata compensated for low productivity and death of infected plants by producing large numbers of tillers.     

The value of timothy (Phleum pratense L.) in ryegrass — timothy mixtures managed to simulate intensive grazing

The performance of timothy in mixtures with perennial ryegrass was assessed under a simulated intensive grazing management over two harvest years in 1974–75. Three seed rates of S23 perennial ryegrass were factorially combined with three rates of Scots timothy and compared with pure stands of each grass. All were sown with Huia white clover. When cut six times at monthly intervals and with an annual N input of 350 kg ha^?1, there were no significant differences in total DM production in either year. The 2-year mean DM yield for the nine mixtures and six pure swards was 9·77 t ha^?1 (range 9·34–10·16). Compared with the pure ryegrass swards, in both years the ryegrass-timothy mixtures produced earlier spring growth but were significantly lower yielding at the second cut. Over the first five cuts the proptortion of timothy in the three mixtures with 22·4 kg ha^?1 ryegrass seed averaged 26% in the first year and 37% in the second. Corresponding calculated mean DM yields of timothy were 2·75 and 3·00 t ha^?1. It is concluded that an early timothy variety is capable of competing with a late-heading perennial ryegrass in frequently cut swards managed to simulate intensive grazing. The strong development of timothy in the dry summer of 1975 suggests that in mixtures of late perennial ryegrass varieties, an early variety of timothy should be beneficial for its spring growth in grazed swards.     

FERTILE TILLER PRODUCTION AND SEED YIELD IN MEADOW FESCUE (FESTUCA PRATENSIS L.)

The effect of defoliation and date of nitrogen application in the spring to seed crops of meadow fescue, S215 and S53, were compared in two harvest years, in drills spaced at 18 in. S215 tillers were generally more robust than S53 tillers, and S215 produced a higher number of fertile shoots as the result of a higher conversion rate of over wintering to fertile tillers. N applied early and absence of defoliation gave the highest ratio of seed heads to over wintering tillers.
Seed- and straw-yields were depressed by April and May defoliations; May cutting had a particularly drastic effect on both varieties. S53 yields in all treatments were generally inferior to those of the hay variety, S215.
Head lengths and spikelet numbers were higher and 1000-seed weight lower in S53 than in S215.     

THE EFFECT OF GROWTH-REGULATING WEEDKILLERS ON THE MORPHOLOGY OF GRASSES

A series of experiments were carried out in 1954 and 1955 designed to investigate the effects of MCPA and 2,4-D applied at different stages of growth to grasses grown for seed. Single, spaced plants were sprayed at the rate of 1·5 lb. a.e. per acre with 2,4-D amine in 1954 and with 2,4-D amine and ester and MCPA ester in 1955. The development of the grass growing point was used as a criterion of the developmental stages at which the plants were sprayed.
Some treatments caused abnormal types of growth. S.23 perennial ryegrass and S.143 cocksfoot developed ear malformations, S.53 meadow fescue had leaf and ear malformations, while S.51 timothy showed leaf, stem and ear abnormalities.
Leaf malformations developed only after spraying when the growing points showed vegetative primordia while ear abnormalities usually appeared after treatment at most early stages of development. Stem malformation in timothy occurred after spraying at all stages in 1954, but only at the vegetative stages in 1955.
The percentage of abnormal ears in a random sample of tillers in all but one case fell as the growing point developed and most damage occurred after treatment when the apex was at the vegetative stage. Normally, treatment when glume ridges were visible on the apex produced no significant difference between the number of abnormal ears in the treated and untreated plots.
The results suggest that the chemicals act on that part of the apex which is just beginning its development but is not yet visible as a protuberance on the apex.
The type of abnormalities occurring in grasses as a result of spraying with growth regulating weedkillers is similar to those found in cereals after similar treatment.
Provided that the chemicals are applied in the 4 to 5 weeks prior to ear emergence the amount of malformation occurring in the grasses will be small.