A comparison of spike and spikelet survival in mainstem and tillers of barley

Initiation and abortion of spikelet primordia on primary tillers and mainstem were compared for plants of spring barley, Maris Mink, sown on two sowing dates during 1983, in plots at the Scottish Crop Research Institute (SCRI) and at Long Ashton Research Station (LARS). Sowing date had no effect on the number of spikelets initiated on either mainstem or surviving tillers and many more spikelets were initiated on equivalent stems at LARS than at SCRI but more spikelets aborted at the former site. The final number of spikelets produced was eventually similar. Spikelet abortion was complete by anthesis and coincided with the period of rapid stem and rachis elongation and with the time over which tiller death occurred. Despite the fact that plants at SCRI produced fewer fertile tillers they outyielded those at LARS because much heavier grains were produced on the mainstem and tiller 1 of plants at the Scottish site. An interval of 2 wk between sowing had only small effects on the components of yield.     

Population differentiation in the tiller architecture of Microstegium vimineum (Poaceae) in relation to habitat

Grass tillers grow by addition of modular units known as phytomers. Differences in phytomer organ size produce subindividual variation with potential adaptive value. Here, patterns in organ mass along tillers in the invasive annual Microstegium vimineum are related to habitat and tiller architecture. In an earlier study, seed families were collected from two populations: one from a sunny, woodland edge and the other from a shady understory in New Jersey, USA. Plants from these seeds were grown in a greenhouse. Phytomers along primary tillers were divided into culms, leaves, and cleistogamous or chasmogamous spikelets and seeds, dried and weighed. These data were used to examine the quantitative genetics of subindividual variation among families and populations, and the relationship of tiller fitness (based on total seed mass) to the mean or subindividual variance of phytomer traits. Phytomer position along a tiller was the major determinant of organ mass. Leaf mass increased from basal to upper nodes; cleistogamous reproductive mass decreased from upper to lower nodes. Phytomer organs were heaviest in the population from the sunny habitat. Family explained < 18% of variation in organ mass. Tiller fitness was positively correlated with mean culm, and leaf mass, but negatively correlated with coefficients of variation. Field‐collected tillers showed evidence of selection for increased leaf mass. Subindividual variation in M. vimineum is mostly due to phytomer position along a tiller and the prevailing light environment. Differentiation between sunny and shady populations suggests selection favors heavier phytomer leaves and culms, especially in the shady understory where this species is most invasive.     

Insights on the role of tillering in salt tolerance of spring wheat from detillering

Tillering is reduced by salinity, with the primary and secondary tillers being more affected than is the mainstem. To understand the importance of tillering in the salt tolerance of wheat plants, two contrasting genotypes of spring wheat (Triticum aestivum L.) were grown in a greenhouse under saline or non-saline conditions and were subjected to five progressive levels of detillering. Regardless of the genotype and salinity, shoot dry weight, seed yield and seed number per plant were all significantly decreased in the treatments where only one or two tillers per plant remained compared with the untouched treatment (more than three tillers), whereas these same variables per tiller tended to be increased on a per tiller (mainstem or substem tiller) basis. The increased seed yield per tiller observed with tiller reduction may be attributed to the enhanced seed number within the spikelet. Under saline conditions, the reductions in shoot dry weight, seed yield and seed number per plant for the salt-tolerant genotype Kharchia were of a greater magnitude in the treatments where only one or two tillers per plant were present compared with the untouched treatment, whereas the magnitude of this reduction in the salt-sensitive genotype Sakha 61 was decreased.     

草地早熟禾种子产量构成因素的品种差异

对草地早熟禾 2 8个品种种子产量及性状进行了研究 .结果表明 ,2 8个品种间种子产量存在极显著差异 (P <0 .0 1) ,最高的品种为Balin和Monoply ,其产量达 6 4 3.4～ 6 5 0 .2kg·hm-2 ,为最低品种Elysee(90 .1kg·hm-2 )的 7.1倍 .生产性状相关分析表明 ,生殖枝数 /株、生殖枝高、花序长及小穗数 /生殖枝 4个性状为影响早熟禾种子产量的主要因素 .依据对种子产量及上述 4个性状测定值的聚类分析 ,可将 2 8个品种的种子生产性能分为极强、很强、强、中等、差和极差 6个类群 ,其中Monoply等 6品种为极强类群 ;Baronie、Balin和Sydsport、Entopper分别为很强和强类群 ;Nublue等 12品种为中等类群 ;Broadway等 4品种和Ram1、Elysee分别为差和极差类群     

Production and dormancy of wild oat (Avena fatua) seed from plants grown under soil waterstress

In 1974 wild oat plants grown from very dormant seed types fA, fB, fC originating from a single field, and in 1976 plants of type fB derived from another location were waterstressed from the time when the spikelets were just beginning to emerge until the seed was fully ripe. The seed production and dormancy of the seed were determined. Waterstress reduced the number of viable seeds per plant by 42% in 1974 and 49% in 1976. The number of viable seed produced on the tillers of the plants were reduced to a greater extent by waterstress than the number produced on their main stems. Some of the differences between the number of viable seeds produced by the three wild oat types were significant. In 1974, 78% of seed from stressed plants was dormant as compared with 90% of that from unstressed plants. Under a given soil moisture, the dormancy of the three types differed little, although seed dormancy was less in tiller seed than in main stem seed. In 1976 seed from stressed plants was 80% dormant, immediately after collection, whereas that from unstressed plants was totally dormant. Storage for 6 months at 25°C decreased these percentages to 17% and 90% and at 5°C to 72% and 98% respectively. Buried in soil immediately after collection, 66% of viable seed from stressed plants gave seedlings in the first autumn after burial as compared with 4% of seed from unstressed plants. Most of the seed from waterstressed plants gave rise to seedlings in the first autumn, and seed from non-stressed plants in the second spring. The α-amylase content was four times greater in seeds from stressed compared with non-stressed plants.     

Frost damage and winter nitrogen uptake by the grass Poa pratensis L.: consequences for vegetative versus reproductive growth

Frost damage can decrease nitrogen uptake by grasses over winter, and it can also decrease biomass production over the following growing season. However, it is not clear to what extent reduced nitrogen uptake over winter decreases grass production, or whether is it merely a symptom of root damage. We examined the growth response of the grass Poa pratensis L. (Kentucky bluegrass) to variation in the timing of freezing and nitrogen availability over winter in London, Ontario, Canada. All tillers were transplanted into untreated soil in early spring, and at peak seed maturation, root, shoot, and reproductive biomass were measured. There was an interaction between freezing and increased winter nitrogen availability, whereby nitrogen addition increased tiller biomass under ambient temperatures, but decreased tiller biomass in combination with a late winter freeze. The nitrogen response of ambient temperature tillers occurred primarily via increased seed production, whereas for frozen tillers seed production was generally absent. Our results support the hypothesis that nitrogen uptake over winter can increase growing season productivity in P. pratensis, but also demonstrate that increased nitrogen availability increases tiller vulnerability to frost. These results have important implications for grass responses to the alteration of soil freezing dynamics with climate change.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE)

Individual tillers of timothy ( Phleum pratense ) growing from seed in pots were labelled and their life history studied over a period of two years.
The leaf axil position of the first-produced tiller depended on its time of origin. A minimum of five leaves on the parent axis was necessary before its appearance, and no tiller emerged from the axil of any of the three terminal leaves of a shoot.
Ear-bearing capacity was high in early-formed tillers, but decreased progressively with later dates of origin despite favourable day length; all tillers arising after the end of July failed to produce inflorescences. A greater proportion of tillers on the main stem formed ears than secondary tillers of the same age. Time of ear emergence varied only within a few days for all tillers arising before the end of April. The number of leaves formed up to ear emergence varied linearly with date of origin and tiller position, the lowest recorded number being six.
Dry weight and length of inflorescence as well as seed yield per tiller varied linearly with the date of origin of tillers. Primary tillers generally had longer and heavier inflorescences and produced more seed per stem than secondary tillers of the same age.
Tillers arising at all times were found to have a limited span of life, not normally exceeding just over a year. The mean age of ear-bearing tillers varied linearly with their date of origin. Death-rates of all tillers generally increased with their age. Relative death-rates of fertile and vegetative tillers rose rapidly in the autumn but decreased during winter and early spring. Vegetative tillers which elongated in the year of formation survived for a shorter time than equivalent tillers in which stem elongation was delayed until the following year.
The perennial habit of grasses is discussed in relation to tiller survival. Successive new formation and death of tillers keep the plant in a condition of continuous dynamic change.     

Rates of Leaf and Tiller Production in Young Spaced Perennial Ryegrass Plants in Relation to Soil Temperature and Solar Radiation

In an experiment designed to investigate the rate of leaf appearanceand tiller production in young spaced plants of three clonesof perennial ryegrass grown in the field, it was found thatthe rate of leaf appearance per tiller increased linearly withmean soil temperature up to approx. 14 °C. The rate of productionof tillers in relation to the rate of leaf appearance (sitefilling) appeared to be virtually independent of weather conditions.In plants which were adequately established, but still relativelysmall, site filling was equal to or exceeded the theoreticalsteady state; all the tiller buds which were being formed weredeveloping into visible tillers. Thus the relative rate of tillerproduction was controlled by the rate of leaf appearance. Inlarger plants site filling was less complete, and site redundancieswere probably caused by within-plant competition for light atthe tiller bases. Lolium perenne L., perennial ryegrass, tillering, leaf production, solar radiation, soil temperature     

The patterns of growth, assimilation of 14CO2 and distribution of 14C-assimilate within vegetative plants of Loliumperenne at low and high density

The patterns of growth, assimilation of ¹⁴CO₂ and distribution of ¹⁴C-labelled assimilate were followed for 12 wk from sowing in individual plants of Lolium perenne grown in miniswards at either low (500 plants m^-2) or high (5000 plants m^-2) density. At the latter density, plants were characterised by a 50% reduction in RGR, by the production of fewer tillers, especially second- and third-order tillers, and by a reduction in mean tiller weight. All the green and senescing leaves of each tiller assimilated ¹⁴CO₂ and the overall assimilatory capacity of a tiller was directly related to its dry weight. At both densities the plant consisted of a main shoot and established tillers with comparable assimilatory activities and a range of developing tillers that assimilated relatively small amounts of ¹⁴CO₂. As each successive primary tiller developed it was supplied with assimilate from the main shoot and the degree of support was inversely proportional to the dry weight of the tiller. At both densities it was concluded that the first primary tiller could be regarded as an independent assimilatory unit when it reached a dry weight of about 25 mg even though some import of main shoot assimilate continued until the tiller was double this weight. The supply of assimilate to the root system was greatly reduced at both densities compared with previous observations on plants grown singly.     

The strategy of the wheat plant in acclimating growth and grain production to nitrogen availability

Two cultivars of spring wheat (Triticum aestivum L.) were grown to maturity in hydroponic cultures. Nitrogen accumulation was controlled by daily growth-limiting additions of nitrate together with all other nutrients in excess. Six different curves of N accumulation were used, with the same relative changes from day to day, but with different amplitudes. These curves were obtained by using the same mathematic formula of the N accumulation curves but varying the value of initial N content. The total amount of nitrogen added varied from 20 mg plant(-1) to 65 mg plant(-1). Plant bioproductivity showed a linear response to accumulated N. The number of grains per plant increased linearly with increased N availability whereas grain weights were essentially unaffected. Grain N concentrations and N content varied slightly, with highest values generally at the lower N availability levels. The quantitatively most important response to increased N availability was an increased number of earbearing tillers per plant. This varied from 0.1 tiller plant(-1) at maturity when given 20 mg N plant(-1), up to about 2 tillers plant(-1) when given 65 mg N plant(-1). Not all tillers that were initiated developed ears. The reduction of tillers seems to be one important mechanism in adapting plant productivity to N availability. Other individual characters influenced by N availability were straw height and the number of spikelets per spike. The two cultivars behaved in a qualitatively similar manner over the range of N availability even though they quantitatively differed in grain size, N concentrations and yield.     

Main stem sink manipulation in wheat : effects on nitrogen allocation to tillers

The role of main stem (MS) sink size on N use by field-grown soft red winter wheat (Triticum aestivum L. cv Hart) was determined. At Feeke's growth stage 8 (last leaf just visible), 100 micromoles of 99 atom% ¹⁵N-ammonium was injected into the lower MS. At anthesis, MS sink size was adjusted by removal of 0, 33, 66, and 100% of the MS spikelets; tiller spikes were left intact. The MS and tiller average kernel size was unaffected by MS sink manipulations. The MS kernel N concentration increased when MS spikelets were removed. Tiller kernel N concentrations were unaffected except when the entire MS reproductive sink was removed, which caused an increase in tiller kernel N concentration. Net losses of MS vegetative N during grain fill were similar for all treatments except for plants lacking MS spikelets, which mobilized 30% less N from the MS. Labeled N was predominately (>90%) associated with the insoluble reduced N fraction of plant tissues at anthesis. Allocation of labeled N to tillers was not proportional to reduction in MS sink size. These results indicate that the reproductive sink on an individual culm has first priority for vegetative N mobilized during grain fill even when sink demand is reduced substantially.     

Tillering in grain sorghum over a wide range of population densities: identification of a common hierarchy for tiller emergence,leaf area development and fertility

Most studies of tiller development have not related the physiological and morphological features of each culm to its subsequent fertility. This introduces problems when trying to account for the effects of tillering on yield in crop models. The objective of this study was to detect the most likely early determinants of tiller fertility in sorghum by identifying hierarchies for emergence, fertility and grain number of tillers over a wide range of assimilate availabilities. Emergence, phenology, leaf area development and dry weight partitioning were quantified weekly for individual tillers and main culms of tillering and uniculm plants grown at one of four densities, from two to 16 plants m(-2). For a given plant in any given density, the same tiller hierarchy applied for emergence of tillers, fertility of the emerged tillers and their subsequent grain number. These results were observed over a range of tiller fertility rates (from 7 to 91%), fertile tiller number per plant at maturity (from 0.2 to 4.7), and tiller contribution to grain yield (from 5 to 78 %). Tiller emergence was most probably related to assimilate supply and light quality. Development, fertility and contribution to yield of a specific tiller were highly dependent on growing conditions at the time of tiller emergence, particularly via early leaf area development of the tiller, which affected its subsequent leaf area accumulation. Assimilate availability in the main culm at the time of tiller emergence was the most likely early determinant of subsequent tiller fertility in this study.     

A Comparison of Main-stem and Tiller Growth in Barley; Apical Development and Leaf-unfolding Rates

Information is presented on the hierarchical relationship betweenthe stems of Proctor barley grown under controlled conditions.Rates of production of foliar and floral primordia and ratesof leaf unfolding were found to follow the order main-stem >primary tillers > secondary tillers. Apical dome size wasinitially substantially greater for the main-stem than for thetillers due principally to the slower transition of the latterto the reproductive condition; by the time that double ridgesare formed on the tiller apices apical dome sizes are similarto that of the main-stem. Transition to the reproductive conditionoccurs at about the same time for the primary tillers. Thisnear-synchrony results in a smaller number of leaves borne onsuccessive tillers, since these have had less time to initiatefoliar primordia, and a measure of synchrony in awn appearance,since rates of car development are similar for these stems. The nature of the hierarchical relationship between main-stemand tillers is discussed.     

Flowering Phenology and Gender Variation in Pennisetum typhoides

Because of the modular structure of pearl millet (an annual grass crop, Poaceae), different tillers of a plant share the same genotype but are subjected to different environmental conditions during their maturation. This allows investigation of the effects of tiller flowering phenology on allocation to resource-producing photosynthetic biomass, sexual functions, and thus tiller gender. All tillers of plants of two families collected from individual maternal plants (represented by 33 and 31 plants each) were analyzed. In both families, allocation to aboveground vegetative biomass decreased as flowering was delayed. On average, late-flowering tillers were 65% smaller than the first ones to flower. The proportion of biomass allocated to reproduction significantly increased with the flowering rank of the tillers, suggesting that translocations of assimilates occurred between early- and late-flowering tillers. In both families, late-flowering tillers produced significantly fewer pollen grains per stamen than early-flowering ones, and female reproductive allocation (expressed as seed mass per tiller) was also affected by flowering phenology. Tillers became increasingly female as flowering phenology progressed. This gender variation is possibly adaptive because pollination efficiency is maximized by plant height. Natural selection may favor a shift toward femaleness to maximize reproductive fitness in small, late-developing tillers.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE L.): IV. THE EFFECT OF NITROGEN, PHOSPHORUS AND POTASSIUM SUPPLY ON GROWTH DURING THE FIRST YEAR

In a sand-culture experiment lasting 21 weeks plants of timothy grown from seed were maintained at all combinations of three levels of nitrogen, phosphorus and potassium. The number and weight of tillers, leaves and ears, and the weight of roots, were determined at the end of the experiment, and for some of the treatments at intervals of 4 weeks. All three nutrients had significant effects, especially N which at its lowest level of supply tended to mask the effect of the other two elements. Potassium influenced tiller numbers least, especially those of primary tillers, but in the presence of high concentrations of N and P it had a large effect on leaf area and dry weight. Relative growth and net assimilation rates responded to varying nutrient supply only in the early stages of growth, so that in general variations in dry weight were associated with nutrient effects on leaf area.     

恢复措施对退化草地羊草种群有性生殖数量特征的影响

在封育和翻耙处理的退化羊草草地上,对羊草种群有性生殖的数量特征进行了比较研究．结果表明,翻耙样地羊草种群密度下降显著,生殖蘖密度变化不显著,生殖蘖穗长、节间长、生殖生长比率、穗节数、小穗数、小花数、饱满籽实数和结实率都有显著增长,籽实产量、净籽实重和千粒重分别是封育样地的2．68、3．49和1．3倍．在翻耙样地中,羊草种群发育有复小穗的生殖蘖比封育样地多16％,生殖蘖上复小穗的数量也显著增加,生殖蘖以相对少的小穗,生产了较多的小花和更多的籽实来提高种群的籽实产量．两个样地中羊草种群的生物量生殖分配在种群对生殖蘖RA1、生殖蘖对穗序RA4、穗序对籽实RA6 3个层次上无显著差异,而种群对穗序RA2、生殖蘖对籽实RA5的生殖分配差异显著,种群对籽实RA3的生殖分配差异极显著．     

Comparison of early development of three grasses: Lolium perenne, Agrostis stolonifera and Poa pratensis

BACKGROUND AND AIMS: To improve the management of grass communities, early plant development was compared in three species with contrasting growth forms, a caespitose (Lolium perenne), a rhizomatous (Poa pratensis) and a caespitose-stoloniferous species (Agrostis stolonifera). METHODS: Isolated seedlings were grown in a glasshouse without trophic constraints for 37 d (761 degrees Cd). The appearance of leaves and their location on tillers were recorded. Leaf appearance rate (LAR) on the tillers and site-filling were calculated. Tillering was modelled based on the assumption that tiller number increases with the number of leaves produced on the seedling main stem. Above- and below-ground parts were harvested to compare biomass. KEY RESULTS: Lolium perenne and A. stolonifera expressed similar bunch-type developments. However, root biomass was approx. 30 % lower in A. stolonifera than in L. perenne. Poa pratensis was rhizomatous. Nevertheless, the ratio of above-ground : below-ground biomass of P. pratensis was similar to that of L. perenne. LAR was approximately equal to 0.30 leaf d(-1) in L. perenne, and on the main stem and first primary tillers of A. stolonifera. LAR on the other tillers of A. stolonifera was 30 % higher than on L. perenne. For P. pratensis, LAR was 30 % lower than on L. perenne, but the interval between the appearance of two successive shoots from rhizomes was 30 % higher than the interval between two successive leaf stages on the main stem. Above-ground parts of P. pratensis first grew slower than in the other species to the benefit of the rhizomes, whose development enhanced tiller production. CONCLUSIONS: Lolium perenne had the fastest tiller production at the earliest stages of seedling development. Agrostis stolonifera and P. pratensis compensated almost completely for the delay due to higher LAR on tillers or ramets compared with L. perenne. This study provides a basis for modelling plant development.     

A modular approach to biomass allocation in an invasive annual (Microstegium vimineum; Poaceae)

The invasive grass Microstegium vimineum grows in low light beneath the canopy of eastern forests in North America by reiteration of modules (phytomers) along a tiller. Basal phytomers are vegetative; terminal phytomers produce a raceme of chasmogamous (CH) spikelets plus an axillary raceme of cleistogamous (CL) spikelets. Additional subterminal phytomers with CL racemes mature basipetally. Allocation to culms, leaves, and CH and CL within phytomers was examined in relation to light conditions for a population in New Jersey, USA. Plants were reared in a greenhouse from seed families of parents in deep shade (2-8% full sun) or sunny, edge habitats. Primary tillers were subdivided into phytomers, dried, and weighed. Tillers from field habitats were similarly treated. For vegetative and subterminal phytomers, allocation to leaves and CH was greatest for the shady habitat. CL allocation decreased from terminal to reproductively immature subterminal phytomers. CH and CL mass was positively correlated with leaf mass, suggesting that reproduction is determined by available photosynthate. CH mass showed a genetic correlation with leaf mass. Developmental plasticity in modular allocation allows Microstegium to maximize fitness when conditions are favorable (e.g., high light along forest edges) by continual maturation of CL caryopses on axillary racemes.     

Effects of seed treatment with a plant growth regulator on growth and tillering in spring barley (Hordeum distichum) cv. Triumph

Doses of the growth retardant, Terpal, were applied to seeds of spring barley. Germination was delayed as the concentration and duration of exposure to Terpal increased. Tiller bud outgrowth was promoted in treated seedlings and at maturity the total tiller production and number of ear-bearing tillers per plant was increased by the Terpal treatment. The yield of the main shoot was similar in both control and treated plants but the mean ear weight of the tillers was reduced in the latter, and thus there was no overall effect of the treatment on yield. Observations on vegetative plants showed that Terpal also reduced the growth of successive main shoot leaves and the elongation and dry weight of the seminal root system. The results are discussed in terms of an early modification in the distribution of resources during establishment that favour the outgrowth of tiller buds at the expense of the initial development of the main shoot and root system. It is proposed that seed treatment is a viable alternative to foliar spraying particularly in arid regions.     

Effect of spikelet removal on the whole plant senescence of rice

The rice (Oryza sativa L.) cultivar IET 1444 showed a nonsequential mode of senescence as evident from the decline in chlorophyll and protein of the flag and second leaves at the senescent stage. Removal of 50,75 and 100 % spikelets from the panicle of rice plant or emasculation of the panicle by hot water treatment induced the development of secondary branch from the axil of second leaf but 25 % removal had no effect. Similarly, removal of 75 and 100 per cent spikelets from the panicle of secondary branch induced tertiary branch development, while 25 and 50 per cent removal had no such effect. Similar treatments on tertiary branch had no effect on further branch production. The pattern of leaf senescence of the untreated (control) main tiller, secondary and tertiary branches was identical, i. e. nonsequential, which could be changed into the sequential type only by the development of additional sinks (i. e. side branch). The leaf area and the seed number of secondary and tertiary branches were gradually reduced and reached a critical value in the tertiary branch. The removal of spikelets or emasculation delayed leaf senescence of the main tiller and the secondary and tertiary branches. Also the longevity of the whole plant could be increased by 40 d i. e., up to the senescence of the tertiary branch. Both leaves and reproductive parts control side branch production, which, in turn, controls the longevity of the whole rice plant.