A Study of Floret Development in Wheat (Triticum aestivum L.)

Plants of wheat (Triticum aestlvum L.) cv. Aotea were grownat high or low nitrogen levels and in a natural photoperiodor continuous light. Starting 17–21 days from the double-ridgestage, eight plants from each treatment were sampled every 3days until anthesis, and the two basal, the sixth, and the terminalspikelets were sectioned longitudinally. A developmental scorewas assigned to each floret and rates of development calculated.Continuous light hastened development but reduced the numberof spikelets per ear, while high nitrogen delayed developmentbut increased spikelet numbers. The number of florets initiatedin each spikelet varied within narrow limits, but grain settingdepended strongly on spikelet position and on treatment. Althoughflorets were initiated in acropetal succession, the rate ofdevelopment tended to increase up to floret 4 but then declinedmarkedly. As a result grain setting was confined to basal floretpositions, although the two basal spikelets developed so slowlythat they contributed relatively little to grain yield. Distalflorets degenerated almost simultaneously at or before ear emergence,but those in intermediate positions continued to develop untilafter fertilization in the lower florets. It is argued thatthe spikelet is an integrated system in which correlative mechanismsplay a part throughout the development of the florets.     

The Effect of Population Density on Floret Initiation, Development and Abortion in Winter Wheat

There is little information in the literature concerning thephysiological basis of the relationship between plant populationdensity and kernel number in winter wheat (Triticum aestivumL.). Thus, two experiments were conducted to evaluate this relationship.Expt 1, involving three population densities, was carried outnear Taian, China in 1982 and in Expt 2, two densities wereevaluated near Lexington, Kentucky in 1986. Plants were sampled every 2 d in the spring, main stem spikeswere dissected and florets were scored according to a 10-stagescale of development. The rate of primordia initiation increasedas density increased until the point at which primordia numberswere equal in all treatments. After this point, an increasein density reduced the primordia initiation rate. In both experimentsincreasing density reduced the total number of floret primordiainitiated and the number of kernels per spike. In Expt 1 theeffect of density on kernel number per spike was accounted forapproximately equally by the effect of density on number ofprimordia initiated and floret abortion. In Expt 2, however,floret abortion was influenced much less by density and accountedfor only 7 % of the variation in kernel number per spike. Thekey result was that the effect of density was determined earlyin floral development. The data suggest that yield losses athigh densities may be determined too early in development tobe offset by N applications at the terminal spikelet stage. Triticum aestivum L., spike development, spikelet development, seeding rate     

Effects of exogenous hormones on floret development and grain setin wheat

At specific stages during floret development, solutions of IAA,GA₃, zeatin and ABA were injected into the leaf sheath around theyoung spike of wheat (Triticum aestivum L.) to study theregulating effects of exogenous hormones on floret development. Zeatin promotedfloret development and significantly increased the number of fertile florets aswell as grain set, especially at the stage of anther-lobe formation. Zeatinalsoincreased the sugar concentrations in spikes at anthesis. In contrast, IAA,GA₃ and ABA inhibited floret development, with different patternsforeach of the hormones. IAA inhibited the development of the whole spike and allflorets in the spikelets such that grain loss occurred in all positions in thespikelets. GA₃ increased the number of fertile florets per spike,butdecreased grain set of the third floret in each spikelet, especially whenapplied at terminal spikelet formation. ABA inhibited floret development, anddecreased the number of fertile florets and grain set at almost all developmentstages, except at anther-lobe formation. The inhibitory effect of ABA wasmainlyon the first and third florets in each spikelet.     

Studies on the Physiology of Flowering of Timothy (Phleum pratense L.) II. Influence of Daylength and Temperature on Size of the inflorescence

Inflorescence length in timothy increases when the photoperiodis reduced from 24 to 14 hours of light; it is also increasedby a reduction in ambient temperature from 75° to 55°F. There is a linear relation between total floret number andear length. Both factors affect ear length by influencing therate of growth of the spike between spikelet initiation andear emergence; this implies an effect on either the number ofprimary spikelet initials or the number of florets producedby branching, or both. Experiments with Lolium temulentum, wheredaylength and temperature influenced initiation and ear developmentin a way similar to that observed in timothy, suggest that thesefactors affect the number of florets at each primary initial.The interrelations of internal and external factors and theirinfluence on inflorescence size in the grasses is discussed.     

Relationships Between Grain Weight and the Size of Floret Cavity in the Wheat Spike

The volume of the floret cavity at different floret positionsalong the spike and within a single spikelet was estimated in10 Triticum aestivum and three T. durum lines by injecting floretcavities with liquid silicone rubber which solidified thereafter.Highly significant correlation coefficients, ranging from 0.40to 0.76 were found between floret volume and grain weight; inmost lines the basal florets had a higher correlation than theterminal ones. The relationships between floret volume and grainweight were characterized by an intercept of about 30–60per cent of the mean grain weight (heavy-grained lines havinga larger intercept) and a slope of about 1 mg µI^–1.Differences in grain size and shape, both within spike as wellas among lines, closely reflected the variation in the sizeof floret cavity. The data support the hypothesis that grainweight is partly determined by the volume of the floret cavity. Grain weight, grain volume, floret cavity     

Reproductive Development in Lolium temulentum L.: Spike Morphogenesis and Grain Set Limitations

Reproductive development in cereals is not easy to investigatebecause their quantitative response to environmental factorsmakes it difficult to synchronize the plants. In this paper,one of our aims was to assess whether Lolium temulentum strainCeres, a qualitative long-day grass, could serve as a modelof reproductive development for cereals. The morphological patternsfrom floral transition to seed set were studied. A flowering scale was established to evaluate developmentalrate during spike morphogenesis. Apex growth was found to increaseaccording to biphasic kinetics; double ridge appearance markedthe beginning of an exponential phase. Developmental progression and apical growth rate were both increasedby giving repeated long days. In contrast, the final numberof lateral spikelets (20–25) could not be manipulatedafter the beginning of long-day treatment. When plants were kept in continuous light from the beginningof induction, double ridges appeared on the fifth 24 h cycle.Spikelet initiation began in the upper mid-part of the spike,and then extended acro- and basipetally. The phase of spikeletinitiation lasted 6 d, with l?5 to 1?9 spikelets being produceddaily. Within each spikelet, florets were initiated at an averagerate of l?3 primordia per day and developed acropetally. Thefirst signs of apical site degeneration were observed in themost developed upper spikelets just before heading. Ear emergenceoccurred between the 20th and 25th cycles of continuous light;anthesis was observed 6 or 7 d later. The proportion of floretssetting grain averaged about 40%. Grains were produced mainlyin the lower spikelets while the upper mid-part of the inflorescenceshowed a much lower fertility rate. Complex developmental gradients described in this paper suggestthat L. temulentum could serve as a model of reproductive developmentin cereals, with the added advantage of flowering in responseto a single long-day. Key words: Lolium temulentum L., spike morphogenesis, spikelet number, floret number, grain set     

Effect of Photoperiod and Chlormequat on Apical Development and Growth in a Spring Wheat (Triticum aestivum) Cultivar

Spring wheat, cv. Timmo, was grown under three photoperiod regimes(16, 13 and 11 h) with and without treatment with the plantgrowth regulator chlormequat (applied at the glume primordiumstage of apical development) and the relationships between apicaldevelopment, primordium initiation and growth stage examined The effects of photoperiod were generally similar to those reportedfrom other studies; shorter photoperiod slowed the rate of apicaldevelopment, increased the duration of the primordium initiationphases and reduced the rate of primordium initiation. The finalnumber of spikelets was increased, but there was no effect onnumber of floret primordia per spikelet The number of tillersproduced was also higher in the shorter photoperiods. Chlormequattreatment had a similar effect to imposing short-days: floweringwas delayed and tiller production increased There were strong correlations between certain development eventsand the phasing of primordium initiation and growth stages andthese were not affected by photoperiod or chlormequat treatments.For example, the end of spikelet primordium initiation, i.e.terminal spikelet (TS) formation, coincided with the floret-stamenprimordium stage (of the most advanced spikelet) and the endof floret primordium initiation with the stigma tic branchesand hairs on ovary wall elongating stage. Similarly, rapid stemextension growth always started at TS formation while spikeextension and spike growth commenced at TS formation and thestigmatic branches stage, respectively. Tiller production alsoceased at TS formation, when rapid stem growth started Although the timing of the phases of primordium initiation andcertain growth events were linked to apical development, therate of apical development did not determine either the rateof spikelet primordium initiation or the rates of stem and eargrowth. However, there was a strong relationship between rateof development and rate of floret primordium initiation. Therewas also a strong relationship between spike length and apicaldevelopment stage Triticum aestivum, spring wheat, photoperiod, chlormequat, apical development, primordium initiation, stem and spike growth     

Differential Effects of the Parental Photothermal Environment on Development of Dormancy in Caryopses of Aegilops Kotschyi

Aegilops Kotschyi Boiss. plants produce three florets per spikelet(of which the terminal floret is commonly sterile) and exhibitphysiological heterocarpy in the two caryopses which developin each spikelet. Germinability of the basal caryopsis, butnot of the upper one, was subject to environmental influencesto which the parent plant was exposed during its development.Basal caryopses produced by plants grown at low temperature,or in 16-h photoperiods, were more dormant than when producedby plants grown at higher temperature, or in 8-h photo-periods.Germinability of the upper caryopses was equally high in allcases and independent of the parental environment. The photoperiodeffects on germinability were exerted after anthesis. Matureweight of both basal and upper caryopses was higher when producedon plants grown at low temperature, or transferred from 16-hto 8-h photopenods at emergence of the flag leaf     

The Vascular System of the Spikelet in Wheat (Triticum aestivum)

The lower three florets in a spikelet of wheat cv. Aotea werefound to be supplied by the principal vascular bundles of therachilla, while the system of more distal florets consistedof subvascular elements derived from the vascular cylinder formedat the disc of insertion of these florets. This pattern appearedto be the same in all spikelets irrespective of position, apartfrom the terminal one, nor was it altered by raising the supplyof nitrogen. The apparent constancy of the vascular system iscontrasted with considerable variability in the number of grain-bearingflorets depending on genotype and environment. If a floret isconnected directly with the main vascular system and if assimilatesare not limiting, then competition for assimilates does notappear to be the main factor preventing grain formation.     

Autophagy regulated by day length determines the number of fertile florets in wheat

SUMMARY: The wheat spikelet meristem differentiates into up to 12 floret primordia, but many of them fail to reach the fertile floret stage at anthesis. We combined microarray, biochemical and anatomical studies to investigate floret development in wheat plants grown in the field under short or long days (short days extended with low-fluence light) after all the spikelets had already differentiated. Long days accelerated spike and floret development and greening, and the expression of genes involved in photosynthesis, photoprotection and carbohydrate metabolism. These changes started while the spike was in the light-depleted environment created by the surrounding leaf sheaths. Cell division ceased in the tissues of distal florets, which interrupted their normal developmental progression and initiated autophagy, thus decreasing the number of fertile florets at anthesis. A massive decrease in the expression of genes involved in cell proliferation, a decrease in soluble carbohydrate levels, and an increase in the expression of genes involved in programmed cell death accompanied anatomical signs of cell death, and these effects were stronger under long days. We propose a model in which developmentally generated sugar starvation triggers floret autophagy, and long days intensify these processes due to the increased carbohydrate consumption caused by the accelerated plant development.     

Heterochronic development of the floret meristem determines grain number per spikelet in diploid, tetraploid and hexaploid wheats

Background and Aims

The inflorescence of grass species such as wheat, rice and maize consists of a unique reproductive structure called the spikelet, which is comprised of one, a few, or several florets (individual flowers). When reproductive growth is initiated, the inflorescence meristem differentiates a spikelet meristem as a lateral branch; the spikelet meristem then produces a floret meristem as a lateral branch. Interestingly, in wheat, the number of fertile florets per spikelet is associated with ploidy level: one or two florets in diploid, two or three in tetraploid, and more than three in hexaploid wheats. The objective of this study was to identify the mechanisms that regulate the architecture of the inflorescence in wheat and its relationship to ploidy level.

Methods

The floral anatomy of diploid (Triticum monococcum), tetraploid (T. turgidum ssp. durum) and hexaploid (T. aestivum) wheat species were investigated by light and scanning electron microscopy to describe floret development and to clarify the timing of the initiation of the floret primordia. In situ hybridization analysis using Wknox1, a wheat knotted1 orthologue, was performed to determine the patterning of meristem formation in the inflorescence.

Key Results

The recessive natural mutation of tetraploid (T. turgidum ssp. turgidum) wheat, branching head (bh), which produces branched inflorescences, was used to demonstrate the utility of Wknox1 as a molecular marker for meristematic tissue. Then an analysis of Wknox1 expression was performed in diploid, tetraploid and hexaploid wheats and heterochronic development of the floret meristems was found among these wheat species.

Conclusions

It is shown that the difference in the number of floret primordia in diploid, tetraploid and hexaploid wheats is caused by the heterochronic initiation of floret meristem development from the spikelet meristem.Key words: Triticum, wheat, inflorescence, spikelet, floret, meristem, heterochrony, heterochronic development, knotted1, polyploidy     

Senescence-induced expression of cytokinin reverses pistil abortion during maize flower development

Maize is a monoecious species that produces imperfect (unisexual), highly derived flowers called florets. Within the spikelet, the basic repeating unit of the maize inflorescence, the spikelet meristem gives rise to an upper and a lower floret. Although initially bisexual, floret unisexuality is established through selective organ elimination. In addition, the lower floret of each ear spikelet is aborted early in its development, leaving the upper floret to mature as the only pistillate floret. Expression from the cytokinin-synthesizing isopentenyl transferase (IPT) enzyme under the control of the Arabidopsis senescence-inducible promoter SAG (senescence associated gene)12 was observed during early maize floret development. Moreover, the lower floret was rescued from abortion, resulting in two functional florets per spikelet. The pistil in each floret was fertile, but the spikelet produced just one kernel composed of a fused endosperm with two viable embryos. The two embryos were genetically distinct, indicating that they had arisen from independent fertilization events. These results suggest that cytokinin can determine pistil cell fate during maize floret development.     

Interactions Among Number of Spikelets, Number of Grains and Grain Weight in the Spikes of Wheat (Triticum aestivum L.)

In a field experiment, comprising four spring wheat cultivars,the frequency and final weight of the grains developing fromeach individual floret were determined in intact spikes andin spikes of which up to nine spikelets had been removed. Theextent of damage caused by the cutting procedure was estimated. Characteristic distributions of the frequencies and weightsof the individual grains were found for each cultivar. Removalof spikelets resulted, in most cases, in a small increase inthe number of grains and in a considerable increase in the weightof the grains of the remaining spikelets. These increases compensatedonly partially, and differently in the different cultivars,for the loss of the removed spikelets. Defoliation at the timeof earing caused a subsequent reduction in grain yield of intactspikes but no reduction in the yield of spikes from which ninespikelets had been removed. The removal of the upper floretsin each spikelet resulted in a certain increase in the weightof the two basal grains. It is concluded that an increase in the number of spikeletsper spike may reduce grain weight but will nevertheless contributeto yield. The number of grains per spikelet is cultivar dependentbut not causally associated with grain weight. Grain set indistal florets is expected to add rather small grains to thespike's yield. Under conditions of limited supplies it may causea reduction in the weight of the basal grains. Any increasein grain weight is anticipated to contribute to grain yieldand is not liable to affect spikelets per spike or grains perspikelet. Wheat cultivars, Triticum aestivum, growth of inflorescence, grain yield, spikelet number     

Responses of Leaf and Tiller Emergence and Primordium Initiation in Wheat and Barley to Interchanged Photoperiod

The influence of constant (9, 13 and 19 h) and reciprocally-interchangedphotoperiods [at terminal spikelet (TS) or triple mound (TM)]on leaf, tiller and primordium development were examined usingphotoperiod-responsive cultivars of spring wheat, ‘UQ189’ and spring barley, ‘Arapiles’. In bothspecies, constant longer photoperiod reduced the duration fromsowing (S) to double ridge (DR), as expected. However, photoperiodsensitivity was not restricted to this mainly vegetative phase.There was also a marked increase or reduction in the durationof reproductive phases between TS/TM and heading (H) when plantswere transferred to shorter or longer photoperiods respectively,compared with controls. These responses were largely independentof the photoperiod during previous phases although minor effectsof the previous photoperiod were observed. For both species,the time course of leaf emergence was linear, or bi-linear,depending on the final leaf number on the main stem. The rateof leaf emergence was faster for the first six to eight leavesthan for the leaves appearing subsequently. The rate of emergenceof early-formed leaves was independent of photoperiod whereasthe rate of emergence of later leaves varied with photoperiod.Photoperiod also affected the dynamics of tillering. The rateof leaf primordium initiation was little affected by variationin photoperiod, but the rate of spikelet initiation increasedwith increases in photoperiod. The rates of leaf and spikeletprimordium initiation were both substantially higher in barleythan in wheat. The fact that the reproductive phase from TS/TMto H was largely independent of the duration of the previousphase provides evidence that this phase might be geneticallymanipulated to increase the time for floret development andhence grain number. Copyright 2000 Annals of Botany Company Photoperiod, rate of development and leaf emergence, tillering, wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.)     

Wheat floret survival as related to pre-anthesis spike growth

Further improvements to wheat yield potential will be essential to meet future food demand. As yield is related to the number of fertile florets and grains, an understanding of the basis of their generation is instrumental to raising yield. Based on (i) a strong positive association between the number of fertile florets or grains and spike dry weight at anthesis; and (ii) the finding that floret death occurs when spikes grow at maximum rate, it was always assumed that floret survival depends on the growth of the spike. However, this assumption was recently questioned, suggesting that assimilates diverted to the spike do not determine the number of florets and grains and that the onset of floret death may instead be a developmental process that is not associated with spike growth. In this study, the relationships between the fate of floret primordia and spike growth from six independent experiments that included different growing conditions (greenhouse/field experiments, growing seasons, photoperiod/shading treatments during the floret primordia phase) and diverse cultivar types (winter/spring, semi-dwarf/standard-height, photoperiod sensitive/insensitive) were re-analysed together. Onset of floret death was associated with the beginning of spike growth at the maximum rate in c. 80% of the cases analysed; and the rate of floret death (the main determinant of floret survival) showed a negative quantitative relationship with spike weight at anthesis. As floret death and survival were shown to be linked to pre-anthesis spike growth, the strategy of focusing on traits associated with pre-anthesis spike growth when breeding to increase wheat yield potential further is valuable.     

The Influence of Genes for Vernalization Response on Development and Growth in Wheat

Studies were made of the influence of genes for vernalizationresponse on the growth and development of four near-isogeniclines of bread wheat (Triticum aestivum L.). The duration from sowing of flower initiation, terminal spikeletformation and ear emergence all increased with increasing vernalizationresponse. There was a close positive relationship between thedays from sowing to flower initiation and from sowing to earemergence, indicating that the duration of either phase of developmentis a useful measure of relative vernalization when daylengthdoes not limit the rate of development. Total spikelet number per ear and the duration of spikelet initationincreased with increasing vernalization response and there wasa correspondingly higher rate of spikelet initiation in thetwo lines with stronger vernalization response. Most of the differences in growth between the lines were associatedwith diferences in development caused by the vrn genes. Maximumtotal above-ground dry matter and total leaf area per plantincreased with increasing vernalization repsonse but relativegrowth rate and leaf area per plant were not significantly differentbetween the lines. There were no differences in net assimilationrate between the four lines until 40 d from sowing; thereafterit decreased, with the greatest decrease in the line with thestrongest vernalization response. Flower initiation, terminal spikelet formation, spikelet initiation, ear emergence, growth rate     

糖蜜草（Melinis minutiflora Beauv．）幼穗分化发育．及花和果实形态的研究

本文对糖密草（ＭｅｌｉｎｉｓｍｉｎｕｔｉｆｌｏｒａＢｅａｕｖ．）的幼穗分化发育及花和果实的形态作了研究,将幼穗分化发育过程划分为以下九个时期：第一苞原基形成期;第一次枝梗原基形成期;第二、三次枝梗原基形成期;小穗及颖花原基形成期;雌、雄蕊原基形成期;花粉母细胞形成期;花粉母细胞减数分裂期;花粉充实期;花粉成熟期。全过程历时约需４２ｄ．从抽穗到颖果成熟约需５０ｄ。糖蜜草的花序为圆锥花序。每花序有可育花２０００—３０００朵．小穗是由小穗轴、内外颖片、不育花外稃和小花构成。小花包括有内外稃各一片、一鳞被、雄蕊三枚和一枚雌蕊,颖果千粒重为９１ｍｇ。     

Soluble Carbohydrates and Floret Fertility in Wheat in Relation to Population Density Stress

Wheat, variety Sonalika, was grown at different densities inboth field and pots during the winter seasons 1983–84and 1984–85. Grain yield pattern of the spikelets of themain shoot inflorescence was similar in both the field and pots,the spikelets in the middle part of the ear contributed mostand yield per spikelet decreased progressively towards the apexand the base. Increased population density decreased yield ofgrain, primarily by decreasing spikelet number and the fertilityof florets. High population density accelerated growth of thespike for some time during the pre-anthesis period and the solublecarbohydrate concentration was also higher under these conditions.During both anthesis and post-anthesis, the soluble carbohydrateconcentration and the growth of the spike declined much fasterin the high-density population. High density also decreasedthe floret fertility and growth in dry weight in all the spikelets,but it was more severe on the basal spikelets, resulting incomplete sterility of the florets at these nodes. The solublecarbohydrate concentration of these slow-growing, sterile, basalspikelets was found to be higher in comparison to that of fertilespikelets in the middle and top positions within the ear. Soluble carbohydrates, spike, spikelet, fertility, grain number     

Floral Development in Sonja White Clover (Trifolium repens) a Papilionoid Legume

Floral development in Sonja white clover was examined usingscanning electron microscopy. Florets and bracts were foundto arise from common primordia initiated as protuberances fromthe apical meristematic area of the inflorescence. The patternof floret initiation on the inflorescence was acropetal, theoldest florets resting basally. Floral organ initiation withineach floret was acropetal, petals being initiated before stamens.Floret development was zygomorphic, each whorl of floral organsdeveloping unidirectionally from the abaxial side. There wasfound to be overlapping in the timing of initiation and developmentof these organs. Antesepalous stamens were found initially tooutgrow their antepetalous counterparts. Early petal developmentwas synpetalous. Eglandular hairs were found basally on thecalyx cup and on the pedicel. Procumbent hairs were found tobe more numerous and randomly distributed on the abaxial surfacesof the mature calyx cup. Trifolium repens L., Sonja cultivar, white clover, scanning electron microscopy, floral development, inflorescence