The effect of defoliation practice in Western Australia on tiller development of annual ryegrass (Lolium rigidum) and Italian ryegrass (Lolium multiflorum) and its association with forage quality

The effect of defoliation on the vegetative, early reproductive and inflorescence stages of tiller development, changes in the dry‐matter yield of leaf, stem and inflorescence and the associated changes in forage quality was determined on plants of annual ryegrass (Lolium rigidum Gaud.) and Italian ryegrass (L. multiflorum Lam.). The field study comprised seventy‐two plots of 1 m × 2 m, sown with one annual ryegrass and seven Italian ryegrass cultivars with a range of heading dates from early to late; defoliation commenced 6 weeks after germination. During the vegetative stage of growth, plots were defoliated when the tillers had three fully expanded leaves (three‐leaf stage). During the early reproductive stage of growth, to simulate a cut for silage, plots were defoliated 6–7 weeks after 0·10 of the tillers displayed nodal development. The subsequent regrowth was defoliated every 3 weeks. Assessments of changes in tiller density, yield and quality were made in the growth cycle that followed three contrasting cutting treatments during the winter–spring period (from 10 July). In treatment 1, this growth cycle (following closing‐up before a subsequent conservation cut) commenced on 7 August following two defoliations each taken when the tillers were at the three‐leaf stage. In treatment 2, the growth cycle commenced on 16 October following: for early‐maturing cultivars, two cuts at the three‐leaf stage, a cut for silage and an additional regrowth cut; for medium‐maturing cultivars three cuts at the three‐leaf stage and a cut for silage; and late‐maturing cultivars, five cuts at the three‐leaf stage. In treatment 3, defoliation up to 16 October was as for treatment 2, but the growth cycle studied started on 27 November following two additional regrowth cuts for early‐ and medium‐maturing cultivars and cut for silage for the late‐maturing cultivars. Tiller development for all cultivars was classified into three stages; vegetative, early reproductive and inflorescence. In treatment 1, in vitro dry‐matter digestibility (IVDMD) and crude protein (CP) content were negatively associated with maturation of tillers. IVDMD ranged from 0·85 to 0·60 and CP ranged from 200 to less than 100 g kg^–1 dry matter (DM) during the vegetative and inflorescence stages respectively. This large reduction in forage quality was due to an increase in the proportion of stem, inflorescence and dead material, combined with a reduction in the IVDMD and CP content of the stem. A high level of forage quality was retained for longer with later‐maturing cultivars, and/or when vegetative tillers were initiated from the defoliation of early reproductive tillers (treatments 2 and 3). However, 15 weeks after the closing‐up date in treatment 1, defoliation significantly reduced the density of inflorescences with means (±pooled s.e_m.) of 1560, 1178 and 299 ± 108 tillers m^–2, and DM yield of inflorescence with means of 3·0, 0·6 and 0·1 ± 0·15 t ha^–1 for treatments 1, 2 and 3 respectively. This study supports the recommendation that annual and Italian ryegrass cultivars should be classified according to maturity date based on the onset of inflorescence emergence, and that the judicious defoliation of early reproductive tillers can be used to promote the initiation of new vegetative tillers which in turn will retain forage quality for longer.     

TILLERING IN GRASSES—ITS SIGNIFICANCE AND CONTROL*

Tillering is examined in relation to the morphology and habit of growth of the perennial herbage grasses; particular reference is made to the effects of cutting and grazing. The importance of tillering in establishment and regeneration or perennation of a grass sward is considered; greater importance is attached to the role of tillers in the regrowth of swards cut for conservation at the reproductive stage of growth. Amounts of regrowth are related to both the number and size of vegetative tillers present at the base of reproductive tillers at the time of cutting. The probable inhibition of the replacement tiller buds by the developing inflorescence is suggested as a reason for the poor growth of grass swards in the favourable environment of midsummer. The physiological mechanisms which control tillering during reproductive development are examined. Spring and winter wheat (Triticum aestivum), Lolium temulentum and Phalaris tuberosa are used as physiological tools to demonstrate that tillering is restricted during reproductive development, and that stem extension may be more important than changes at the stem apex in controlling this restriction of tillering at the base of the inflorescence-bearing stem. It appears from results of the application of a range of growth regulators that substances derived from the meri-stematic centres, in either the elongating stem or the terminal meristem, indirectly control the metabolic activity of the lateral buds from which new tillers are derived. The degree of apical dominance is shown to depend on the light intensity in which the plants are grown. Finally, the physiological results are used to suggest reasons for the commonly found apparent inability of heavily fertilized perennial ryegrass (Lolium perenne) swards to regrow after cutting at the inflorescence-emergence stage of growth.     

The relationship between tiller appearance in spring and contribution to dry-matter yield in perennial ryegrass (Lolium perenne L.) cultivars differing in heading date

The relative contribution of tillers present in April and those appearing in consecutive periods in spring was assessed for perennial ryegrass cultivars in the three maturity groups (early, intermediate‐ and late‐heading). Each group was represented by two diploid and one tetraploid cultivar each in plots in their third (2000) and fourth (2001) harvest years in three replicated blocks receiving an average of 325 kg N ha^?1 and cut seven (in 2001) or eight (in 2000) times annually. ‘Main’ tillers and their daughters were marked with colour‐coded PVC‐covered wire loops in early April as were daughters which appeared in consecutive periods between harvests, the loop colour identifying the period of origin of the tiller. Tillers were harvested at cutting height (5 cm) before the plots were harvested and the herbage from tillers with the same colour code bulked per plot. Tillers were identified retrospectively as ‘reproductive’ if they had been decapitated at the previous harvest. Dry‐matter yield was higher in the early than late‐heading cultivars in April and early May but this was reversed in harvests in late May and June. The early heading group had a lower lamina content than the late‐heading group during reproduction growth, both due to the reproductive tillers (mainly those which overwintered) having a lower leaf content and to their being fewer and smaller vegetative tillers during the reproductive phase than for the late‐heading group. Turnover of tillers was high in spring due to decapitation of reproductive tillers and rapid post‐flowering tillering. This was particularly pronounced in the early heading group which also had slightly more tillers marked in April which were subsequently decapitated than in the other maturity groups, i.e. 0·56 compared with 0·44 for the late‐maturing group. Mean ratios of rate of death: rate of tillering for 3 years (1999–2001) for the early and late‐heading groups were 0·8 and 0·4, respectively, for April–May and 1·1 and 2·4, respectively, for June indicating the different patterns in tiller turnover for the two extreme maturity groups. Information on tiller origin and contribution to yield can be used to refine tiller‐based grass growth models.     

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.     

The physiology of tiller death in grasses. 2. Causes of tiller death in a grass sward

The causes of tiller death in a 2-year-old perennial ryegrass sward were examined between April and August 1977. Physiological causes accounted for most tiller deaths and grazing by slugs and rodents was more important than the damage caused by stem-boring larvae. Tillers which died were mainly small and vegetative, although some flowering tillers died prematurely. Low nutrient status delayed but did not prevent tiller death. Using ¹⁴CO₂ it was shown that small tillers fixed relatively less radiocarbon than did larger tillers and they did not receive much support for their carbon economy. Selective defoliation showed that in April defoliated tillers imported radiocarbon from undefoliated tillers but that in July at anthesis an undefoliated reproductive tiller retained most of the carbon it fixed, despite its vascular association with defoliated tillers. It appears that much of the tiller death during the period April-August is due to the failure of the more favourably placed tillers to support other tillers which are heavily shaded.     

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.     

The effects of nutrient supply on flowering and seed production in Poa annua

The effect of nutrient concentration on the reproductive development and seed yield of Poa annua was examined in a sand culture experiment. The nutrient concentration during the initial vegetative stage did not affect the time taken for double ridge formation by the main shoot but did influence the subsequent development of the inflorescence as did the post-initiation level of nutrients. At low nutrient levels flowering was inhibited in some individuals but at the higher concentrations inflorescence emergence was hastened, inflorescence size was increased and, in particular, the number of spikelets and hence the number of seeds per inflorescence was greatly increased. The mean weight of 100 seeds was unaffected by the nutrient concentration. The number of reproductive tillers per plant was increased by high nutrient supply but the proportion of dry weight allocated to root development was reduced.     

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.     

Physio-Morphological Characters of F1 Hybrids of Rice (Oryza sativa L.) in Japonica-Indica Crosses: II. Heterosis for leaf area and dry matter accumulation

Summary

A pot experiment was conducted to investigate the heterosis for morphological characters and to examine the relationship among some characters at active tillering, flowering and dough ripe stages of 12 Fj hybrids from crosses between japonica and indica rice. Heterosis for plant height, number of tillers, green leaf area and dry matter accumulation per plant was positive at all stages. The intensity of heterosis was higher at the active tillering stage for number of tillers, leaf area and dry matter accumulation per plant. A significant positive relationship was found between tiller number and leaf area per plant at active tillering and flowering stages. Significant positive relationships between leaf area and dry matter accumulation, and between tiller number and dry matter accumulation per plant were observed at all stages. A significant positive relationship was found between leaf area per plant at an early stage and dry matter accumulation per plant at a later stage, suggesting that early development of leaf area is an important factor for higher dry matter accumulation in 1 hybrids. Although heterosis for percent dead leaf blade was positive at the flowering stage due to early leaf senescence in some Fx hybrids, a larger leaf area was also found in Fx hybrids at this stage and leaves of Fx hybrids remained green longer compared to parent cultivars up to the dough ripe stage.     

SEASONAL TILLER POPULATIONS OF GRASS AND GRASSJCLOVER SWARDS WITH AND WITHOUT IRRIGATION

The number of tillers per unit area of pure swards of S23 and S24 perennial ryegrass, S37 cocksfoot, S48 timothy and of a S23 perennial ryegrass/white clover sward, were examined at intervals of 4 weeks during a period of two years. The monthly appearance of new tillers and the death of tillers of S23 and S24 ryegrass and S48 timothy were followed for 15 months. The studies were made with and without irrigation.
The number of tillers varied with season through the year. Tiller numbers increased rapidly in late winter or early spring, and then declined until midsummer. Numbers rose again between June and August. This seasonal pattern of tiller numbers was most marked in ryegrass and timothy and least in cocksfoot. The effect of irrigation on tiller numbers was variable, and usually small in relation to the overall seasonal fluctuations in tiller numbers, but dry soil conditions could delay an increase in tiller numbers between June and August. The relationship between seasonal fluctuations in tiller number and root growth of swards is discussed.     

Natural diversity in vegetative and reproductive investments of perennial ryegrass is shaped by the climate at the place of origin

In forage grasses, vegetative and reproductive investments are major determinants of yield and persistence. A survey of the diversity of vegetative and reproductive investment traits was carried out on 213 perennial ryegrass genotypes, representing 51 natural European accessions. Plants were phenotyped for traits related to leaf elongation, tillering, reproductive investment and heading date, at two locations with markedly different climates. Strong genetic effects for all traits were found. Interactions between genotype and location were moderate. Plants showed stronger spring leaf growth, lower numbers of tillers and higher reproductive investments at Lusignan (France) than at Melle (Belgium). Plant growth rate and tillering capacity were genetically nearly independent suggesting breeding for both traits simultaneously and independently should be possible. A high genetic diversity was observed for all traits. This diversity was structured in three main clusters. One cluster comprised early‐flowering genotypes with high reproductive investments and high spring growth rates. The remaining genotypes fell into two clusters based on differences in tillering capacity. Clear links were found between traits and the climatic conditions of the region of origin of the accessions. Autumn growth rate was positively correlated with solar radiation at the region of origin in October and February.     

Effect of heading date of perennial ryegrass cultivars on tillering and tiller development in spring and summer

Three cultivars (two diploid and one tetraploid) in each of three maturity groups (early, intermediate and late) of perennial ryegrass were sown in 10 m² plots, replicated four times, in Northern Ireland in June 1997 in a study of the effect of heading date on tiller development (including initiation to flower) and turnover of tillers produced at specific times in spring in 1998 and 1999. The plots were harvested seven times in each year. Annual dry‐matter production was similar for all groups in each year. In spring and early summer of both years, tiller density of the diploid cultivars was 1·5 times greater than that of the tetraploid cultivars and the mean tiller density over all swards in June was about 0·40 times greater than that in April. Maximum proportions of reproductive tillers in the early, intermediate and late maturity groups, determined from apical dissections, were found in early April, mid‐May and early June, respectively. Although a high proportion of tillers, which were present when annual observations commenced in spring, was decapitated at the first harvest in the early group, the previous population density was maintained by rapid production of new tillers during May, including those from suppressed tiller buds during reproduction. It is concluded that the relationship between heading date and rate of tiller turnover (including flowering) at specified times in spring is important in sward management throughout the early part of the growing season and should be taken into account in tiller‐based grass growth models.     

Is there a tiller morphology ideotype for yield differences in perennial ryegrass (Lolium perenne L.)?

The objectives of this study were to measure the variability in ‘realized’ tiller morphology and identify whether there are emergent generalizations about a tiller ideotype for productivity or survival. Morphological traits during the vegetative growth stage were measured for 2 years in the field for eight perennial ryegrass cultivars: six diploid and two tetraploid cultivars under low and high nitrogen fertilizer (50 and 225 kg N ha^?1 year^?1 respectively). Traits measured were lamina width, length and area; pseudo‐stem length and diameter; an index of tiller shape; tiller dry weight; tiller density; and herbage mass. Almost all the traits differed significantly between cultivars and significant correlations were found between the 2 years. Principal component analysis identified that tiller morphology and dry‐matter yield were independent. No trait associations indicating alternative cultivar‐specific tiller morphologies were detected but cultivars differed in tiller size. The patterns of change between the 2 years indicated a possible morphological trajectory as swards age. The genetic compensatory relationship between tiller size and density had a 1:1 slope, indicative of constant yield. This implies that larger tillers would be more leafy, and higher leafiness of tetraploid over diploid cultivars was confirmed by the leaf:non‐leaf ratio and tiller shape index.     

四川小麦分蘖冗余及理想群体构成研究

为探究四川小麦分蘖冗余及理想群体构成,以5个典型冬小麦品种为材料,在分蘖始期至拔节期每2d剪一次分蘖,设置3个处理[仅保留主茎,剪除所有分蘖(B0);保留主茎和1个大分蘖(B1);保留主茎和2个大分蘖(B2);不剪分蘖(CK)],对成熟期小麦株高、穗部性状和单株产量进行调查分析。结果表明,5个品种间各性状对剪除分蘖的反应表现较一致。单株穗数以B0处理最低(1.00穗),B1、B2和CK处理单株穗数在同一水平(1.96~1.98穗);B0处理下穗粒数(49.5粒)、千粒重(53.2g)、穗长(11.3cm)、有效小穗数(17.7个)较CK显著增加,但株高(86.5cm)和单株产量(2.54g)均最低;B2处理的单株产量(4.27g)介于B1(4.39g)和CK(4.13g)之间,但与二者差异均不显著;B1处理下株高(88.1cm)适中,主茎和分蘖的穗长(11.0、10.6cm)增加,有效小穗数(17.8、16.4个)增多,最终主茎和分蘖的穗粒数(47.4、39.5粒)显著高于B2(44.2、34.7粒)和CK(43.8、36.0粒),主茎和分蘖的千粒重(50.7、52.5g)略高于B2(50.3、50.9g)和CK(49.6、51.3g),穗数、穗粒数、千粒重协调,单株产量(4.39g)最高。因此认为,保留主茎1个分蘖处理消除了冗余分蘖对资源的浪费,减少了内耗,是四川小麦优质、低耗、高产的最佳茎蘖组合。     

INTER-VARIETAL COMPETITION IN PERENNIAL RYEGRASS SWARDS

Pure cultures and mixtures of equal numbers of plants of Irish. New Zealand and S23 perennial ryegrass were grown in artificially constructed swards under field conditions. They were cut every time they reached grazing height for three successive years. Counts of plant numbers and tiller numbers per plant made each November show that S23 suppressed both plant number and tiller number per plant of Irish ryegrass and to a lesser extent of New Zealand perennial ryegrass. Its own plant number and tiller number per plant were improved in these combinations. New Zealand perennial ryegrass suppressed plant and tiller numbers of Irish perennial ryegrass.
Varying the method of cutting to match the growth of each variety resulted in improved plant survival and tiller number per unit area of Irish perennial ryegrass when cut according to the growth of S23, i.e. late in the spring. The importance of the effect of experimental technique is discussed.     

THE EFFECT OF TIME OF CUTTING ON EAR PRODUCTION AND SEED YIELD IN S.48 TIMOTHY

After uniform defoliation in February, spaced plants of S.48 timothy were cut on one of 12 fortnightly dates, starting at the end of March.
Analysis of the herbage removed at each cut showed an increase in plant and unit tiller weight throughout the season, but a decline in the number of tillers, especially before the beginning of shooting.
The date of ear emergence was not affected by cutting up to 5 May and only slightly delayed in plants cut on 19 May. Plants treated subsequently flowered much later but after a fairly constant interval following defoliation.
Cutting on and after 7 April depressed fertile tiller formation with increasing severity, until only vegetative tillers appeared in the recovery growth following treatment at the beginning of August.
Seed yield per plant declined with the decrease in fertile tiller numbers, but after the early cuts some compensation by production of slightly more seed per ear was recorded.
The results are discussed in terms of the physiological transition of the plant towards flowering.     

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.     

The effect of autumn fungicide application on spring dry matter accumulation in perennial ryegrass cv. S24

Fungicide applied to plots of perennial ryegrass cv. S24 in the autumn of 1981 significantly increased tiller number and total dry matter accumulation in February and March of 1982. However, fungicide application in the autumn of 1982 failed to increase tiller number or dry matter accumulation in the spring of 1983. It is suggested that fungicide application in autumn 1981 prevented the death of tillers caused by fungal attack after snow covered the plots in December 1981.     

SEED PRODUCTION STUDIES ON PERENNIAL RYEGRASS, TIMOTHY AND PRAIRIE GRASS

A study over two years involving tiller identifica tion according to month of origin enabled the contribution of individual tillers of perennial rye grass Grasslands Ruanui, timothy Grasslands Kathu and a local New Zealand strain of prairie grass (Bromus unioloides) to be determined at seed harvest. In addition, the distribution of seed-heads in different arbitrary emergence groups was detemined so as to allow analysis of individual seedhead components within each group. In all species, tillers formed during the first 4 months following an autumn sowing in the first year, and also in the immediate post-harvest period and through the autumn in the second year, made a major contribution to seedhead numbers and seed yield at harvest. These tillers had a lower vegetative mortality rate than those formed in the spring. Primary tillers formed after sowing were highly persistent and became almost exclusively repro ductive. Many vegetative tillers died before har vest, behaving as annual tillers. Spring-formed tillers made a major contribution to the vegetative growth of the plant over the summer and early autumn. These tillers generally died before the following winter. Tillers formed in winter and early spring by ryegrass and prairie grass showed a high mortality, particularly in the second year. In timothy, mortality was highest amongst tillers formed in late spring. The reduced contribution of spring-formed tillers to the seed head population occurred as a result of fewer heads numerically compared with older tillers. In all species, most seedhead components varied according to time of ear emergence. Early emerged heads were generally longer, bore more spikelets and more florets per head and had a greater culm length than late-emerged heads. The major trend in all species was a general depression in head length, spikelet and floret number and culm length with grazing. These effects were in most cases partly or completely overcome by N application. The major effect of applied N was to increase floret number per head.