Tiller dynamics and assimilate partitioning in Lolium perenne with particular reference to flowering

The fate of 100 seedling plants of Lolium perenne L. was studied over a period of 2 years in a field plot. The birth and death of tillers and the production of inflorescences was followed, and the components of seed yield were recorded in detail in the first year. The pattern of distribution of ¹⁴CO₂ assimilated by the main shoot was examined at monthly intervals and during the first flowering season the distribution of ¹⁴C-assimilate from individual leaves and from the inflorescence was also studied. The capacity of individual tillers to assimilate ¹⁴CO₂ prior to flowering and the re-distribution of previously accumulated assimilate during seed growth were also assessed. Plants died at a more or less constant rate with time and only 54 survived to the end of the 2–yr period. First year mortality was associated with severe grazing or cutting but in the second year the death of ungrazed plants was observed. There was great variability in the production of tillers by surviving plants. In both years the number of live tillers per plant increased from July to the end of April with particularly rapid tillering in March and April establishing the maximum value for each year. There was a similar phase of rapid tillering after flowering in July. The number of live tillers per plant declined by 50% during stem elongation and inflorescence emergence and the majority of dead tillers were young secondary (in the first year) and tertiary (in the second year) tillers with a mean age of 40 days. Such tillers had poor assimilatory capacity prior to the onset of death and were not supplied with assimilate from the main shoot. Most of the plants surviving at the end of the experiment flowered in both years and one quarter of the maximum number of live tillers per plant recorded in April of each year produced inflorescences. The earlier a tiller was produced the greater was its chance of flowering and the greater its production of seed. The greater weight of seed produced was associated with the development of more seed-bearing florets per spikelet. There was relatively little export of “C-assimilate from the flowering main shoot, and the lower internodes formed the major sink for post-anthesis assimilate. The growth of seeds appeared to be relatively independent of the leaves for current assimilate. There was some evidence that assimilate accumulated in lower internodes was remobilised and utilised in the growth of seeds and new tillers. Overall, the results confirm the view that the grass plant is a dynamic population of short-lived tillers and indicate that increasing competition for assimilate at flowering exerts a major influence on the production and survival of tillers.     

Tillering in grain sorghum over a wiide range of population densities: modelling dynamics of tiller fertility

The prediction of tillering is poor or absent in existing sorghum crop models even though fertile tillers contribute significantly to grain yield. The objective of this study was to identify general quantitative relationships underpinning tiller dynamics of sorghum for a broad range of assimilate availabilities. Emergence, phenology, leaf area development and fertility of individual main culms and tillers were quantified weekly in plants grown at one of four plant densities ranging from two to 16 plants m(-2). On any given day, a tiller was considered potentially fertile (a posteriori) if its number of leaves continued to increase thereafter. The dynamics of potentially fertile tiller number per plant varied greatly with plant density, but could generally be described by three determinants, stable across plant densities: tiller emergence rate aligned with leaf ligule appearance rate; cessation of tiller emergence occurred at a stable leaf area index; and rate of decrease in potentially fertile tillers was linearly related to the ratio of realized to potential leaf area growth. Realized leaf area growth is the measured increase in leaf area, whereas potential leaf area growth is the estimated increase in leaf area if all potentially fertile tillers were to continue to develop. Procedures to predict this ratio, by estimating realized leaf area per plant from intercepted radiation and potential leaf area per plant from the number and type of developing axes, are presented. While it is suitable for modelling tiller dynamics in grain sorghum, this general framework needs to be validated by testing it in different environments and for other cultivars.     

An appraisal of ear, leaf and stem 14CO2 assimilation, 14C-assimilate distribution and growth in a reproductive seed crop of amenity Lolium perenne

In seed crops of Lolium perenne, yield may be reduced by competition for a limited assimilate supply from sinks other than the ear. This study was undertaken to evaluate the priorities for allocation of assimilate within the crop from all photosynthetic sites on the main reproductive tiller after anthesis. Ear, stem and leaves were fed with ¹⁴CO₂ on two occasions; the assimilatory efficiency of these sources and the magnitude and pattern of ¹⁴C-assimilate export from each was determined. The growth of each part of the main tiller and subtending tillers was also measured. Stem elongation apparently dominated the current assimilate resource and the ear did not become a net importer of assimilate until this process had ceased. Assimilate allocation to the tillers was high throughout. The nature of any competition between these sinks is discussed. When crop growth was contrasted with that in a previous year, environmental factors were implicated as determinants of priority for assimilate allocation to each sink. Sources of carbon for seed filling are also discussed as is the relevance of these findings to seed crop management.     

Cocksfoot (Dactylis glomerata) Grown for Production of Seed

In grasses grown for production of seed under field conditions,it has been found that only a small proportion of tillers produceinflorescences. For a period of 23 weeks from 4 February 1963a study was made of cocksfoot grown for production of seed inan attempt to discern whether the ability of tillers to becomereproductive was related to specific morphological features.Most tillers which initiated spikelets were rooted, and rootsquickly developed on unrooted tillers which had initiated spikelets.It appeared that larger tillers, judged on the basis of dry-weight,were more likely to become fertile than smaller tillers; importantfeatures of tiller size were the length of the stem and theweight of green leaf (lamina) borne by the tiller. The potentialnumber of inflorescences was reduced considerably by the death,in June, of tillers with completely differentiated apices.     

Pattern and Control of Distribution of 14C-assimilates in Reproductive Plants of Lolium multiflorum Lam. var. Westerwoldicum

The distribution of ¹⁴C-assimilates was examined in reproductiveplants of Lolium multiflorum Lam. var. Westerwoldicum (cv. Tama)from which all emerged tillers had been removed, leaving themain tiller with two expanding leaves, one of them the flagleaf, and two expanded leaves. Export of ¹⁴C from the lowerexpanded leaf was mainly to the tiller in its axil, the steminternode below its node and the roots, whereas the upper expandedleaf supplied predominantly the expanding leaves, the ear, steminternodes, roots and the tiller bud in the axil of the lowerleaf. Defoliation and root-pruning showed that expanding leaveswere able to compete successfully for assimilates, probablythrough the production of substances capable of mobilizing supply.Local application of 1-naphthaleneacetic acid (NAA), gibberellicacid (GA₃) and 6-benzylaminopurine (BAP) to small tiller budsshowed that GA³ and BAP promoted bud growth and ¹⁴C accumulation,but that addition of NAA reduced these effects.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE)

All tillers of timothy ( Phleum pratense ) growing in sand culture were labelled at weekly intervals. At each of eighteen dates between June and December a sample of plants was harvested, and dry weight and leaf area were determined for each group of tillers.
Tiller production was continuous throughout the experiment. Only those tillers which appeared before the end of July were able to form inflorescences, but among them the proportion of tillers with ears decreased with increasing lateness of origin.
Dry weight per plant increased rapidly for the first 14 weeks, and then did not change significantly until the end of the experiment. The flowering tillers lost dry weight after seed maturation; vegetative tillers, although smaller in size, continued to increase in number and weight. Dry weight per inflorescences was greatest in the main stem and decreased in subsequent tillers.
Leaf area per plant rose to a maximum soon after ear emergence and then declined, largely because the loss in leaf surface sustained by the flowering tillers was not offset by a corresponding increase in the vegetative tillers. Leaf area ratio decreased at first rapidly but later more gradually, while net assimilation rate was relatively high until maximum leaf area was almost attained, but then fell continuously to negligible values by October. Relative growth rate decreased throughout the experiment.
Initially the growth of the plant was largely determined by the flowering tillers in which, in close resemblance to the cereal plant, leaf area ratio and relative growth rate declined continuously. Tillers which did not flower were successively smaller in size and, because of their later origin, were exposed to less favourable environmental conditions by the time they had increased sufficiently in number to affect the growth of the plant.     

刈割对多年生黑麦草分蘖与叶片生长动态及生产力的影响

对不同刈割强度和时间下 ,混播草地中多年生黑麦草净分蘖数和叶片生长季节动态及其年积累和年产量进行了 2年的比较研究。结果表明 :叶片净生长与净分蘖数季节动态具有相互弥补的特点。刈割不利于多年生黑麦草的分蘖 ,所有刈割处理均不存在分蘖的“超补偿生长”效应 ;净分蘖数随刈割次数的增加而降低 ;高频刈割对分蘖的抑制效应逐年加强 ,低频刈割对分蘖的抑制效应逐年减弱。叶片生长对刈割处理的超补偿效应显著 ;刈割能刺激多年生黑麦草叶片的生长 ,提高年净生产力 ,维持混播草地中黑麦草的组分 ,增强黑麦草的竞争能力。刈割对多年生黑麦草年净生产力的提高主要是通过刺激其叶片的生长来实现的。叶片净生长与净分蘖数的年积累存在显著负相关 (R=- 0 .893,P<0 .0 5)     

冬小麦拔节期不同茎蘖对低温胁迫的反应及抗冻性评价

以小麦济南17和山农8355为材料,在低温胁迫条件下,测定了不同茎蘖功能叶和叶鞘超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)活性及丙二醛(MDA)和可溶性蛋白含量,并利用主成分分析、聚类分析对其抗冻性进行综合评价.结果表明： 低温胁迫条件下,小麦拔节期不同茎蘖功能叶和叶鞘中SOD、POD和CAT活性均不同程度地上升,MDA和可溶性蛋白含量则不同程度地上升或下降.利用主成分分析和聚类分析,将济南17不同茎蘖分为3类：主茎和一级分蘖Ⅰ、Ⅱ属强抗冻蘖组,一级分蘖Ⅲ、Ⅳ和二级分蘖Ⅰp属中度抗冻蘖组,二级分蘖Ⅱp属弱抗冻蘖组;将山农8355不同茎蘖分为3类：主茎和一级分蘖Ⅰ、Ⅱ、Ⅲ属强抗冻蘖组,一级分蘖Ⅳ和二级分蘖Ⅰp属中度抗冻蘖组,二级分蘖Ⅱp属弱抗冻蘖组.表明冬小麦拔节期不同茎蘖存在抗冻性差异,且低位蘖较高位蘖抗冻.
     

The effect of tiller removal and tiller defoliation on competition between the main shoot and tillers of spring barley

In glasshouse and field experiments the source-sink relations of the main shoot of plants of spring barley were modified by tiller removal and tiller defoliation. Decreasing competition by tiller removal promoted the growth of the residual main shoot and its component parts, and the earlier tillers were removed the greater was the effect. Stem dry weight was increased four-fold in the glasshouse by early tiller removal and was doubled in the field experiment. The grain yield of the main shoot ear was increased by 26 – 30% by tiller removal compared with tillering control plants and this was due to larger grains in all spikelet positions. On the other hand increasing competition by regular tiller defoliation had relatively little effect on the growth and development of the main shoot in the glasshouse study, but in the field the main shoot grain yield was reduced by 10% compared with the control. The main effect of tiller defoliation was on the development of tillers. In the glasshouse tillers survived repeated defoliation, continued to be produced, and the majority produced grain but with fewer and smaller grains per ear than in control plants. Tiller growth was supported by the import of assimilate from the main shoot and this was accompanied by an increase in the photosynthetic rate of the main shoot leaves. In the field all defoliated tillers died within 4 wk. These responses are discussed in terms of the physiological interrelations between the main shoot and tillers.     

松嫩平原野大麦无性系分蘖株的年龄结构

野大麦无性系分蘖株由３个龄级组成,为明显的增长型年龄结构、１龄级蘖在无性系生殖生长中占绝对优势。分蘖株龄级越高,其数量增长速率越小,对无性系物质积累的贡献越小,１、２龄级分蘖株数量和生物量均随着无性系丛径和总蘖数的增加而增长,３龄级蘖与总蘖数间呈直线相关。平均单蘖生产力随着龄级增加而下降。生殖蘖平均单蘖重具有相对稳定性。无性系中１龄级蘖平均单蘖重体现出一定的密度调节作用,无性系潜在种群也为明显的增     

Carbon dioxide exchange in relation to sink demand in wheat

Summary In this paper, experiments are described which examine the effect of requirement for assimilates by the ear on the rate of net photosynthesis in leaves of wheat (Triticum aestivum L.). Different levels of requirement were achieved by various levels of sterilization of florets just before anthesis, which resulted in a range of grain numbers per ear, and by inhibiting photosynthesis of the intact ear by 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU). Only the ear and two uppermost leaves of the main shoot were considered, all the lower leaves and tiller leaves being excised when the experimental treatments were imposed. In two experiments, tiller regrowth was permitted during the experimental period, while in a third, new tillers were defoliated regularly.The response of leaf photosynthesis to the level of assimilate requirement by the ear was influenced by the treatment of the vegetative tillers. Thus, the net photosynthesis rate of the flag leaf was decreased by a reduction in grain number, or increased by inhibition of photosynthesis in the ear, only when the vegetative tillers were kept defoliated; when these tillers were allowed to re-grow normally, there was no influence of ear treatment on leaf photosynthesis. Temporal changes in leaf photosynthesis were consistent with this response pattern, i.e., when tillers were defoliated, the initial high rates of photosynthesis persisted for much longer.In the experiment where photosynthesis was influenced by the requirement for assimilate in the ear, the variation occurred through change in stomatal conductance on the abaxial surface of the leaf. This surface has a lesser conductance to CO₂ exchange than the adaxial surface. The implication of this finding to rapid methods of plant screening is discussed.     

Estimation of Potential Tiller Production and Site Usage During Tall Fescue Canopy Development

Tiller appearance in tall fescue (Festuca arundinacea Schreb.)occurs in an orderly, predictable manner with the potentialfor a high degree of synchronization among tillers on a givenplant. Estimates of potential cumulative tiller production (Tmax)are made for synchronous (Tmax_lx = 2^Lx+1 – 1, where Lxis the axil number of the youngest leaf on the main stem whichbears an emerged tiller) and non-synchronous (Tmax₁ = 2^L–NLAT–1,where L is the number of leaves on the main stem and NLAT thenumber of leaves above the youngest primary tiller at its appearance)conditions. A method for determining the degree of synchronizationand an equation for estimating site usage are also presented.Early in seedling development, site usage of a tall fescue populationwith high tillering capacity was near 90%, and tillering wasregulated largely by rate of tiller site formation. As the canopydeveloped the phyllochron (time between successive leaf appearances)and NLAT increased, slowing the rate of tiller production intemporal terms and in relation to leaf appearance, respectively.Beyond 45 d after planting, site usage decreased rapidly, furtherreducing tiller production. High tiller production appearedto be associated with synchronized tiller appearance, with alack of synchrony being associated with decreased site usage.Tillers formed in prophyll axils were less likely to be in synchronywith other tillers and frequently failed to appear. In contrastwith branching in dicotyledons, apical dominance appears toplay a minor role in regulating tillering in tall fescue Tillerproduction initially appears to proceed at near maximum ratesthen is down-regulated during later development by longer phyllochrons,slower rate of tiller elongation and reduced site usage. Festuca arundinaceaSchreb., tall fescue, tiller production, leaf appearance, site usage, leaf elongation rate, synchronization, phyllochron, canopy development     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE L.) III. ABSORPTION AND DISTRIBUTION OF NITROGEN, PHOSPHORUS AND POTASSIUM DURING THE FIRST YEAR OF GROWTH

Absorption of nitrogen, phosphorus and potassium by S48 timothy plants ran parallel to dry-weight increase, at an increasing rate until the autumn, almost completely ceasing in the winter, and increasing again in the spring.
Flowering tillers accumulated nutrients until ear emergence, after which the content per tiller of all nutrients decreased until death of the tillers during the winter.
Non-flowering tillers continued to accumulate nutrients steadily throughout the whole period of the experiment.
These observations are discussed in relation to the perennial habit of the plant.     

Effects of Shading the First Leaf on Growth of Barley Plants: I. Long-term Experiments

Plants of barley were grown under controlled conditions andthe first or second leaves covered with tubular shades thusreducing the light intensity at the leaf surface to low levels.Expansion of the shaded leaves was not prevented, but appearanceof the next leaf but one and all subsequent leaves on the mainstemwas delayed by up to 3 days. Primordia of the first four leaveswere present in the dry grain. Shade treatment delayed slightlythe initiation of the eighth and subsequent leaves and transitionto the double ridge stage at the mainstem apex. Shading the first leaf caused a temporary reduction in the rateof dry-matter increase of plants, but after 14 days the ratewas similar to that of control plants. Smaller effects werefound when the second leaf was shaded. Dry-matter productionfollowed two logarithmic phases in the period prior to awn emergence,and rates for the whole plant and for plant parts were similarfor control and shaded plants. Thus, apart from the initialperturbation, shading had no effect on growth in terms of rateof dry-weight gain. Shade treatment did not affect weight per grain or numbers ofgrain per ear, but over-all yield of grain was significantlyreduced since shading delayed the appearance of tillers andalso reduced the number of tillers bearing grain. The effectof shade was especially marked on tillers originating on primarytillers. Similar qualitative effects on tiller development werefound in an experiment on wheat.     

放牧干扰对根茎冰草无性系种群年龄结构的影响

根茎冰草（Agropyron michnoi)是典型的无性系禾草。在松嫩平原,根茎冰草的分蘖节最老为４龄,可存活５个年度。不同季节的放牧干扰可导致根茎冰草无性系种群分蘖株的年龄结构发生变化,全年放牧样地和夏秋休牧样地均为稳定型年龄结构,两个冬季放牧样地均为增长型年龄结构。１龄分蘖株的生产力和营养繁殖力力均最大。４龄分蘖株均已丧失了营养繁殖力。营养繁殖芽的年龄结构可以更好地预测预报无性系种群和群落的未来。全年放牧可保护稳定型年龄结构,冬季放牧可保持增长型年龄结构,夏秋休牧可使种群发展为增长型年龄结构,两个冬季放牧样地将仍然保持单优势种群落,全年放牧地将仍为混生群落,夏季林牧样地具有向着单优势种群落演替的潜在趋势。     

Developmental ideotype in the context of the dual‐purpose use of triticale,barley and durum wheat

Phenology strongly affects the performance of dual‐purpose (forage plus grain) cereals by defining the rate of leaf and tiller appearance, the duration of the pre‐grazing period and anthesis date. Here, the combined effect of genotypic and environmental variation conditions on anthesis date and the pre‐grazing period in durum wheat, triticale and barley was analysed to define a developmental ideotype for their dual‐purpose use. The extensive variation for the Haun stage reached at the time of apex emergence from the soil surface was associated with variation in the final leaf number, regardless of environment, species and cultivar. The timing of the terminal spikelet stage was the best predictor of apex emergence. The durum wheats tillered poorly, whereas the barleys and the triticales performed comparably in three of the four environments. In general, the higher the final leaf number, the higher the number of leaves and tillers produced by the time of apex emergence, the higher the number of leaves which emerges subsequently. The development of a high number of leaves both before and after apex emergence is advantageous, provided that anthesis is not overly delayed such that flowering and grain fill occur within a period where stressful conditions are likely to occur. Barley developed many leaves and had a low phyllochron, so was identified as the species with the development best suited for dual‐purpose cropping. However, the near simultaneous emergence of the first two tiller apices exposes it to a risk of grazing damage.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE L.)

During the first year of growth of timothy, variations in nitrogen supply affected the number of tillers and leaves, leaf area and dry weight, but had little influence on relative growth and net assimilation rates, except at very low levels of nitrogen. Differentiation and ear emergence occurred slightly earlier at high nitrogen and ear length was increased; there was also an increase in the proportion of fertile tillers among the total number produced after the first few weeks of tillering. The effect of nitrogen supply on the number, dry weight and ear length of early primary tillers was small but became larger in later tillers. Irrespective of the level of nitrogen supply, tillers had particularly high net assimilation rates during the first few weeks following their appearance.     

Photosynthesis and transpiration of the flag leaf in four spring-wheat cultivars

Co₂ exchange and transpiration rates of the flag leaves of four spring wheat (Triticum aestivum L.) cultivars, namely Glenlea, Neepawa, Opal and Kolibri, were compared using infra-red gas-analysis technique. The plants were grown in a controlled environment under an 18-h photoperiod, with day and night temperatures of 20 and 15° C, respectively. The time course of the CO₂-exchange rate (CER) of the flag leaf differed among cultivars. CER began to decrease rapidly some 2 weeks after ear emergence in Glenlea, Neepawa and Kolibri, but only after 4 weeks in Opal. The decline in CER of Glenlea, Neepawa and Opal was continuous throughout the period of grain development whereas in Kolibri CER was maintained at a constant level between the 4th and 6th weeks after ear emergence. The transpiration rates of the flag leaves of the 4 cultivars did not change markedly until 6–7 weeks after ear emergence, indicating that the reduction in CER was not primarily a response to increased stomatal resistance to the diffusion of CO₂. Removing the ear of the main shoot of intact plants failed to depress CER of the subtending flag leaf until 5 weeks after ear removal. Removing the ears of all the tillers of plants in which all but 3 tillers had been removed at ear emergence did not depress CER until 4 weeks after ear emergence, but removal of the ear of the main shoot of plants where all the tillers had been removed at ear emergence reduced the CER of the flag leaf 2 weeks after ear removal. Removal of tillers at ear emergence had a marked effect on the time course of CER and transpiration rates of the flag leaf. Both CER and transpiration rates of a 4-tiller plant were maintained at a higher level throughout ear development as compared to those of a one-tiller plant. The transpiration rate of the flag leaf of Glenlea increased during the later part of the life of the leaf even for one-tiller plants with no ear, indicating that such a stomatal response may be part of the normal course of leaf aging and not a response to a feedback stimulus from the ear.     

Leaf Blade Dimensions of Rice (Oryza sativa L. and Oryza glaberrima Steud.). Relationships between Tillers and the Main Stem

The effects of genotype and environment on the leaf area indexof rice are well documented, but the rules governing leaf areadistribution among main stem and tillers are less well understood.This study investigated the ontogenetic patterns of leaf bladearea and dimensions on the main stem and tillers of three ricecultivars, IAC47, Javaé (Oryza sativa L.) and CG14 (O.glaberrima Steud.) grown in pots without competition among plants.No differences in phyllochron were observed among culms, butleaves on tillers appeared 0.5 to 0.8 phyllochrons earlier thanthe genealogically corresponding leaves on the main stem (systemof Katayama). Cohorts were thus not fully synchronized. Allculms produced their largest leaves soon after panicle initiation(PI), despite differences in tiller age. Leaves that appearedsubsequently were smaller owing to reduced length but not width,which remained constant. Three different hypothetical rulesgoverning the behaviour of culms were tested: (1) tillers behaveas clones of the main stem; (2) cohorts behave uniformly, and(3) behaviour is determined by leaf position (sum of genealogicalleaf and tiller indices). The results were best explained byhypotheses (2) and (3), with leaf blade width and length showingmarkedly different patterns. Copyright 2001 Annals of BotanyCompany Phenology, phyllochron, leaf area, leaf position, leaf cohorts, summed leaf position, leaf blade length and width