THE GROWTH AND DEVELOPMENT OF SOME GRASS SPECIES UNDER COMPETITIVE STRESS

Three experiments were carried out in a semi-controlled glasshouse environment to determine the response of individual plants to competition; (i) between seedlings of different species, (ii) between seedlings derived from seeds of differing sizes within a species, and (iii) between seedlings and established plants.
Differences were detected in seedling competitive ability between species and between plants derived from differing seed sizes. In associations of species of very high and very low competitive ability the differences in competitive ability were accentuated by an increase in plant density.
Established plants of Phalaris coerulescens were less aggressive to seedlings growing in close proximity than were plants of Lolium rigidum and H1 ryegrass. Despite the early death of established swards of L. rigidum , and the consequent removal of competitive stress, there was no rapid increase in the growth and development of seedlings in these swards. Possible mechanisms causing this effect are discussed.
There was evidence of a specific relationship between the species of the established sward and the seedling species, in particular seedlings of HI ryegrass were particularly susceptible to competition from established plants of their own species.
Dry weight per plant, tiller production, rate of leaf appearance and leaf size were ail affected by competition between seedlings; however, tiller production was generally more severely affected than rate of leaf appearance and leaf size.
Dry weight per plant, tiller production, seedling height, number of leaves per tiller and number of seeds per seedling were all affected by competition from established plants.     

The nature of competition between perennial ryegrass and white clover

Perennial ryegrass and white clover were grown in boxes with either no competition, root competition only, shoot competition only, or both root and shoot competition between them, Boxes received either no nitrogen or 200 kg N ha⁻¹ as a split application. The experiment was harvested at 8, 12, 16 and 20 weeks after sowing. Perennial ryegrass was more competitive than white clover throughout the experiment and its overall competitive ability increased with Lime. The effects of root competition were greater than those of shoot competition at the first harvest, with both forms of competition having similar affects at the second harvest, while at the third and final harvests shoot competition had greater effects than root competition. At the later harvests N application increased the overall competitive ability of perennial ryegrass, relative to white clover, mainly owing to the increase in its shoot competitive ability.     

THE GROWTH AND DEVELOPMENT OF SOME GRASS SPECIES UNDER COMPETITIVE STRESS

Two pot experiments were carried out in a semi-controlled environment to examine the response of individual plants to competitive stress in simple systems incorporating varying cutting frequencies and soil–nitrogen levels. A third experiment was set up outdoors to examine the effect of competition on individuals in an association, on the association as a whole, and to extend the controlled environment studies to a more complex model involving micro-plots.
Phalaris coerulescens , although its seedlings possessed a high competitive ability, showed a dramatic decline in competitive ability in later stages of the association, this decline being associated with extensive floral development. In micro–plots the effect of competition was so severe as to prevent the expression of seasonal growth characteristics by P. coerulescens .
Considerable inter-specific differences existed in the response to competition of leaf-area production per tiller during periods of regrowth.
Whilst P. coerulescens remained highly productive and maintained a dense tiller population in monocultures, its tiller regeneration following flowering was severely suppressed in competition with vegetative plants of HI ryegrass or Festuca arundinacea .
The number of heads produced per plant, head length and time of head emergence were all affected by competitive stress, the former response being a reflection of the relative competitive abilities at the time of floral initiation. The response of the latter two characters reflected the decline in competitive ability which occurred during later stages of the growth and development of P. coerulescens .     

THE EFFECT OF CUTTING ON THE SUBSEQUENT GROWTH OF INTRODUCED RIBGRASS (PLANTAGO LANCEOLATA) SEEDLINGS

The ability of ribgrass to germinate and subsequently develop in a sward was investigated as part of a competitive study between this species and perennial ryegrass S23. The experimental model emphasized that seedling development was greatly influenced by the frequency and the intensity of cutting to which the associated grass species was subjected.     

A comparison of root competition and shoot competition between Dactylis glomerata and Holcus lanatus

Dactylis glomerata and Holcus lanatus were grown together in boxes with either full competition, root competition only, shoot competition only or no competition between them. The boxes were either fertilized (150 kg ha^-1N, 100 kg ha^-1 K, 80 kg ha^-1 P) or not fertilized. The experiment was continued for 13 months, and a total of six cuts were taken.
H. lanatus was more competitive than D. glomerata throughout the experiment, though the effect declined after flowering during the second growing season. The effects of root competition were generally much greater than those of shoot competition and tended to increase during the first year. However, the effects of root competition declined, relative to shoot competition, after flowering in the second growing season.
Fertilizer applications slightly increased the root competitive ability of H. lanatus relative to D. glomerata but slightly reduced its shoot competitive ability.     

Potential allelopathic influence of certain pasture weeds

Laboratory and growth chamber research was conducted to determine the potential allelopathy of tissue harvested from certain weeds commonly found in pastures. Results of bioassays indicated that the weeds dogfennel and mayweed chamomile are potentially allelopathic to alfalfa and Italian ryegrass seedlings. Leaf-tissue extracts from the two weed species reduced seedling growth more than stem and root extracts. In most experiments alfalfa seedlings were influenced more than Italian ryegrass seedlings and seedling growth was influenced more than seed germination. Leaf tissue from the weed species mixed into potting soil at a concentration as low as 0.25% (w/w) significantly reduced alfalfa and Italian ryegrass plant development and foliage production. Extract and tissue concentration used in this research were estimated to be similar to concentrations expected to occur within the pasture ecosystem and below concentrations that would result in hypertonicity in the bioassays.     

Effect of season and cutting frequency on root and shoot competition between Festuca pratensis and Dactylis glomerata

Abstract Plant competition strongly affects the species composition of managed grassland. To identify relevant processes, Festuca pratensis (Huds.) and Dactylis glomerata (L.) were grown as monocultures or mixtures in boxes placed in the field for two seasons and subjected to two cutting frequencies. Root and shoot competition effects were separated using soil and aerial partitions. Shoot competition was analysed by measuring the vertical distribution of the leaf area and root competition by analysing the absorption of tracers. Values of relative yield indicated that the two grasses fully competed for the same limiting resources under the experimental conditions. The competitive ability of F. pratensis was lower during both years relative to D. glomerata. This was mainly related to its consistently lower shoot competitive ability, which was associated with less leaf area in the upper layers of the canopy and shorter leaves. Root competitive ability of F. pratensis changed with season. It was similar to that of D. glomerata during spring and autumn, but less during summer. The lower root competitive ability in summer might be due to the lower root activity of F. pratensis, measured as rubidium (Rb) and strontium (Sr) absorption in July, 0·1 and 0·2 m below the ground. Cutting frequency did not greatly influence the relative importance of root and shoot competition during the first growing season. However, under infrequent defoliation, the competitive ability of F. pratensis decreased markedly in full competition during the second growing season. These results suggest that distinct differences in the canopy structure and root activity of different plant species can affect the relative importance of root and shoot competition during the season.     

混播比例对热研2号柱花草和飞机草早期生长竞争能力的影响

为明确热研2号柱花草(Stylosanthes guianensis cv.Reyan No.2)与飞机草(Chromolaena odorata(L.)R.M.KingH.Robinson)生长早期的竞争关系,探索飞机草替代控制的途径和管理策略。设置总株数固定(8株),5种柱花草和飞机草分株数混播比例(柱∶飞=8∶0、6∶2、4∶4、2∶6、0∶8)盆栽试验,就各混播比例下2种植物在浇水频率1次/2 d、生长30 d的生长情况和相对竞争能力进行定量分析。结果表明:随着柱花草混播株数的减少,柱花草和飞机草的相对株高、相对平均生长速度、相对单株总生物量和总生物量分配至根、茎和叶的相对生物量均逐渐降低;但飞机草的相对根长、相对根冠比和根生物量质量分数均逐渐减小、茎生物量质量分数无显著变化、叶生物量质量分数逐渐增大,而柱花草的各对应指标反之。柱花草比例较高时(柱∶飞为6∶2)2种植物的相对产量总和显著大于1,2种植物间无种间竞争现象,4∶4和2∶6时相对产量总和均与1相比无显著差异,2种植物间存在资源竞争;混播群落中柱花草的竞争平衡指数均显著小于0,说明生长早期柱花草的相对竞争能力比飞机草弱。下一步进行生长中期、晚期竞争试验和肥力、刈割等其他因素影响试验来确定柱花草替代控制飞机草的可能性。     

Growth of white clover in eight grass species and the effects of propyzamide, a grass-suppressing herbicide

Seedlings of white clover (cv. Grasslands Huia) were introduced as spaced plants into 3-year-old monoculture plots of eight grass species ( Agrostis capillaris, Agrostis stolonifera, Dactylis glomerata, Festuca rubra, Holcus lanatus, Lolium perenne, Phleum pratense and Poa pratensis ) during June 1984. In mid-April 1985 plots were split for application of propyzamide at the following concentrations: 0, 0·2, 0·4, 0·6 kg a.i. ha^-1.
During 1984 differences between clover seedling growth in the different grass species became apparent within 2 weeks; growth was greatest in F. rubra, P. pratensis and H. lanatus and smallest in D. glomerata, L. perenne and P. pratense. During 1985, when more N fertilizer was given, H. lanatus and D. glomerata, were equally competitive and clover contributed only 16–18% of the total herbage yield of 10·4 t ha^-1 in them, compared with 33–50% of yields ranging from 9·6 to 119 t ha^-1 in the other six species. Propyzamide decreased grass growth in mid-season by more than 50% but there was little overall persistent benefit to clover yield, except for A. stolonifera and P. pratense with 0·4 and 0·6 kg ha_-1 treatments. Dactylis glomerata and perenne were least and H. lanatus, F. rubra and P. pratensis most suppressed by the herbicide.
Possible reasons for the overall large clover contribution from a very sparse seedling population and the relatively small effects of propyzamide are discussed as well as future work required to improve the predictability of effects of grass suppression.     

水稻苗期发根力的QTL和上位性分析

以典型籼粳交(窄叶青8号/京系17)F[sub]1[/sub]花培加倍的DH群体为材料,采取水上栽培方法,考察苗期根系发根力。利用已构建的分子连锁图谱,采用基于混合的线性模型复合区间作图法对水稻苗期发根力进行QTL和上位性分析。在第3染色体的C63-CT125之间检测到1个发根力的主效QTL,同时也检测到影响发根力的5对上位性效应基因座,分别位于第2、3、5、6、7、12染色体上,其中影响根长、根数上位性效应各有两对区间,有一对既影响根长,又影响根数。     

THE RELATIONSHIP BETWEEN COMPETITION AND DEFOLIATION IN PASTURE

Field trials were carried out on a downland permanent pasture at the Grassland Research Institute, Hurley, between 1953 and 1955, to investigate the factors involved in competition in pasture undergoing periodic defoliation. Response to competition was estimated by comparing the yield of species units (either individual plants or close groups of shoots) growing normally in the pasture with the yield of matched units from which the surrounding herbage had been removed. The species studied were cocksfoot, red fescue, creeping bent, plantain, ox-eye daisy and bulbous buttercup. In addition to the estimation of normal competition for environmental resources in pastures of differing height, two special effects upon species due to the presence of surrounding herbage were investigated following unusual results from competition tests in swards under periodic defoliation. First, the influence of surrounding herbage in modifying the growth habit of a species, causing it to grow in a more erect manner, and consequently to suffer more on defoliation at a specific height than in the absence of competition. Secondly, a decreased yield after defoliation, not associated with changes in the growth habit of the species, which was ascribed to an increase in top/root ratio brought about by shading during the preceding period of competitive growth. The effect of selective defoliation of a species in relation to the herbage surrounding it was also studied. Differences in the degree of defoliation of a species and its surrounding herbage had a marked effect upon the degree of competition exerted by the latter upon the former, increasing it when the species was preferentially defoliated and decreasing it when the surrounding herbage was preferentially defoliated. The factors involved in pasture competition under periodic defoliation are listed and discussed.     

Effect of nutrient application and herbicides on root competition between green panic seedlings and a Heteropogon grassland sward

Root competition was studied between seedlings of green panic ( Panicum maximum var. trichoglume ) and Heteropogon contortus in a native grassland sward clipped regularly to a height of 5·7 cm, and to which supplementary irrigation was applied. The influence of fertilizer addition and root competition from the grassland sward on seedling growth during the 42 days following emergence was determined. Root competition was excluded by steel tubes, or eliminated by killing the native sward with herbicides.
Seedling growth was poor where neither root competition was controlled nor fertilizer was applied; at 42 days the seedlings were still less than 6 cm high with three leaves and one tiller. When root competition was controlled, either by steel tubes or herbicide, seedling growth increased significantly. Growth was greatest in the herbicide-treated plots where fertilizer was applied, seedlings at day 42 being 48 cm high with sixty-three leaves and nineteen tillers.
Competition for nutrients appeared to be the primary factor limiting seedling growth and this was largely overcome by fertilizer application. Competition for nutrients appeared to increase the fertilizer requirements of pastures oversown into live swards compared with that of pastures oversown into herbicide-treated swards.     

Root and shoot competition between established grass species and newly sown seedlings during spring growth

Competition between established Dactylis glomerata or Festuca pratensis / Taraxacum officinale swards and seedlings of Trifolium pratense was studied under field conditions in Norway. Root competition from the Festuca pratensis / Taraxacum officinale sward was significantly greater than that from a Dactylis glomerata sward. The balance in root and shoot competition varied between experimental years because of variation in environmental conditions. In the first experimental year, root competition had a greater effect on seedling dry weight than shoot competition, whereas the opposite was found in the second year. However, both root and shoot competition (full competition) in comparison with no competition gave proportionally the same effects in both years, indicating that an alleviation in root competition will be followed by intensified shoot competition. An increasing effect of shoot competition with time on plant dry weight was found. In a second experiment, seedlings of Lolium perenne and Trifolium pratense produced the highest plant dry weights in competition with an established Phleum pratense sward, followed by seedlings of Poa pratensis , Phleum pratense and Trifolium repens in decreasing order. However, no interactions between seedling species and competition treatments were found, indicating no differences in sensitivity between seedling species to competition from the established grass sward. The results are discussed in the context of renovating grassland by direct drilling.     

Effects of defoliation and fertilizers on root competition between Dactylis glomerata and Lolium perenne

Dactylis glomerata and Lolium perenne were grown in single-species stands and in mixed-species stands in boxes for 1 year. Plants were grown in paired rows in each box with full competition below ground, but aerial partitions prevented shoot competition. The rows were cut either frequently (six cuts) or infrequently (three cuts). In single-species stands, frequent defoliation reduced the root competitive ability of D. glomerata plants, when growing with infrequently defoliated plants, though this occurred only if no N fertilizer was applied. Frequent defoliation did not affect the root competitive ability of L. perenne plants, in single-species stands, whether or not N fertilizer was applied. In mixed-species stands, the root competitive ability of D. glomerata was always greater than that of L. perenne, though N fertilizer applications and frequent cutting both reduced the root competitive ability of D. glomerata relative to L. perenne. Mixtures of frequently cut and infrequently cut plants, in unfertilized single-species stands, always yielded more than the mean of the uniformly treated stands (frequently or infrequently cut); they usually yielded more than the highest-yielding uniformly treated stand. Mixtures of frequently and infrequently cut plants, in fertilized single-species stands, always yielded less than the mean of uniformly treated stands. Mixed-species stands usually yielded more than the mean yield of the component species, and sometimes yielded significantly more than the highest yielding component. This effect generally increased with time. Relative yield totals (RYT) were consistently greater than 1·0, in both single-species and mixed-species stands, when no N fertilizer was applied. RYT were greatest (1·2–1·6) when neither N nor P fertilizer was applied, and generally increased with time. The results are discussed in relation to the effects of cutting on root competition and the probable value of mixed-species stands.     

COMPETITION BETWEEN PERENNIAL RYEGRASS AND MEADOW FESCUE UNDER FIELD-PLOT CONDITIONS

Perennial ryegrass and meadow fescue were grown alone and in association in field plots, to gain an idea of the nature and importance of the competitive interactions between them, and their relative contributions to the yield of the newly-established sward.
Experiments over 10 weeks and 9 months both showed that the contribution by fescue to the overall dry-matter yield was severely restricted by perennial ryegrass. There were no deaths of fescue plants during the experimental period (which included an overwintered treatment). The depression in yield was rather a result of a marked decrease in size and weight of the individual plants, brought about by competition for light and nutrients from the faster-growing ryegrass.     

Survival of plants of common sainfoin (Onobrychis viciifolia Scop.) in competition with two companion grass species

A 2-year experiment on competition between sainfoin ( Onobrychis viciifolia Scop.), meadow fescue ( Festuca pratensis ) and tetraploid perennial ryegrass ( Lolium perenne ) was conducted using plants grown in plastic containers outdoors. Root and shoot systems of sainfoin and the grasses were separated in order to investigate full and no competition of root and shoots, at two planting ratios (0·33 grass:0·66 sainfoin and 0·66 grass:0·33 sainfoin). Survival of sainfoin plants was lower at the higher grass:sainfoin ratio. More sainfoin plants died in the winter than during the growing season. Root competition had no effect on survival of sainfoin plants, but shoot competition reduced survival of sainfoin plants during one growing season. Companion grass species had no effect on survival of sainfoin plants, except in the first winter, when fewer sainfoin plants survived when grown with meadow fescue than perennial ryegrass.     

Influence of root competition on compatibility between white clover and perennial ryegrass populations during seedling establishment

Plants of a white clover population, Ac 3785, were grown in binary mixtures with perennial ryegrass cv. Talbot and the 'coexisting' population, Ba 10281, in boxes. Treatments imposed were 'no root competition' and 'full competition', which prevented or allowed the root systems of the competing species to intermingle respectively. Shoot competition was permitted in both treatments. Measurements of dry-matter yield and leaf area of the clover and grasses were made on three occasions following sowing.
The dry-matter yield of both grasses was higher than that of clover at all three harvests and under both treatments, and this yield advantage tended to increase with time. Both grasses yielded more under full competition than where root competition was prevented. Talbot was higher yielding than the coexisting population under both treatments. Clover dry weight and leaf area index (LAI) were considerably higher under the 'no root competition' treatment. There was no effect of companion grass on clover dry weight, but clover LAI was significantly higher with its coexisting population than with Talbot in the 'no root competition' treatment.
The LAI and dry weight data indicate that clover leaf morphology changed in response to the companion grass where root competition was prevented. Thus, the superior aerial compatibility of Ac 3785 with its coexisting grass was limited by below-ground factors. Possible ecological reasons for this effect are discussed.     

Effect of agrochemical applied at sowing on seedling emergence and herbage yield of perennial ryegrass and white clover

Plots of white clover ( Trifolium repens ) and perennial ryegrass ( Lolium perenne ) white clover were sown with and without fungicide seed treatment, seed treatment with Cytozyme (a proprietary growth-promoting agent) and pesticide granules (carbofuran or carbosulfan) mixed with the seed. Sowings were made in April/May at Hurley (1986 and 1987), Dorchester (1987) and Bronydd Mawr (1987). Only treatments containing pesticide significantly improved seedling emergence or herbage yield in the 1986 sowing. In 1987, pesticide greatly increased seedling emergence of ryegrass and clover at Hurley and Dorchester, but had no effect at Bronydd Mawr. At Hurley, herbage yield and the proportion of clover were more than doubled for up to a year, by chemical treatment after sowing in 1986.
Populations of plant parasitic nematodes were recorded at all sites and it is suggested that at Hurley and Dorchester seedlings with nematode damage were unable to survive the dry soil conditions that prevailed. In a subsequent pot experiment, pesticide granules controlled root endoparasitic nematodes and significantly increased seedling emergence of white clover in dry soil but had no effect in moist soil.     

ANALYSES OF YIELD FROM A COMPETITION TRIAL WITH PERENNIAL RYEGRASS

Four varieties of perennial ryegrass, (S24, S321, S23 and Endura), were grown in monoculture and in all possible binary mixtures. Three spatial arrangements of varieties within mixtures were used. Total DM yields were recorded from nine harvests during 1969 for mooocultures and mixtures and for the mixture components. The arrangement of varieties within mixtures had no effect on the total DM yield of the mixtures. Monocultures and mixtures did not differ significantly in yield at any harvest or in annual total, bnt there were differences between monocultures at some harvests. No differences between mixtures were detected. The experiment took the form of a competition diallel in which monocultures corresponded to the parents in a genetic diallel and mixtures to the crosses between these parents. There was a significant interaction hetween the rows (i.e. main variety effects) and the columns (i.e. associated variety effects) of the diallel table only at the first harvest. The change in the competitive ability of the four varieties throughout the season was examined. The relative ranking order of varieties for competitive ability altered as the season progressed.     

THE DIGESTIBILITY AND VOLUNTARY INTAKE OF S22 AND H.1 RYEGRASS, S170 TALL FESCUE, S48 TIMOTHY, S215 MEADOW FESCUE AND GERMINAL COCKSFOOT

Pure swards of S22 Italian ryegrass, H.1 hybrid ryegrass, S170 tall fescue, S48 timothy, S215 meadow fescue and Germinal cocksfoot were each harvested at several stages of maturity during first growth in the spring, and in the subsequent regrowths. Each batch of herbage was fed to sheep in a digestion trial under conditions of voluntary intake. Results from the first growth herbages showed the expected fall in digestibility with increasing maturity. However, the digestibility of S22, H.1, S170 and S48 began to fall well before ear emergence, whereas the digestibility of Germinal fell only slowly up to the time of ear emergence, as had been found previously with S24 and S23 ryegrass and S37 cocksfoot. The rate of fall of digestibility in H.1 and S48 (0.4 units/day) also appeared to be lower than with the perennial ryegrasses and cocksfoot (0.5 units/day). The monthly regrowths within each herbage were of similar digestibility, but the mean digestibility of regrowths differed between herbages, being highest for S215 and lowest for Germinal and S170. The grasses with highest digestibility in the early Spring, in particular S22 ryegrass, were not, however, always the most digestible in the subsequent regrowths. The numbers of sheep fed on each herbage were insufficient to show whether there were significant differences in voluntary intake between different herbage species of the same digestibility. It is suggested that an intake/digestibility relationship, based on data from more animals, should be developed for each herbage species. Only if different relationships are found can it be concluded that factors other than level of digestibility have an effect on intake.