Floret Initiation and Development in Spring Wheat (Triticum aestivum L.)

The number and developmental stages of florets were determinedin each spikelet of the spike in the main shoots of spring wheat.Samples were taken frequently from plants grown in a phytotronand in a nitrogen application field-test. Ten stages of development,from floret initiation until anthesis, were recognized and described. Inter-spikelet variation in the total number of initiated floretswas rather small. However, the number of florets at advancedstages of development, as well as the number of grains, washighest in the central spikelets in which florets initiatedfirst. Floret initiation did not proceed beyond spike emergence,whereafter the distal florets and the spikelet apex degenerated.Grain-set was restricted to florets which had developed at leastto the stage of visible anther lobes at spike emergence. Thenumber of these florets was increased significantly by nitrogenapplication. Wheat, Triticum aestivum L., spikelet, floret, grain set, nitrogen     

The Effect of High Temperature at Different Stages of Ripening on Grain Set, Grain Weight and Grain Dimensions in the Semi-dwarf Wheat 'Banks'

Grain number in the wheat cultivar Banks was reduced by up to11 % with a rise in temperature from 21/16 °C to 30/25 °Cover a 10-d period immediately following first anthesis in general,the upper ‘d’ and ‘c’ florets were moreaffected by high temperature than the basal ‘a’and ‘b’ florets within a spikelet and florets fromthe upper spikelets were more sensitive than those lower onthe ear Grain weight and grain length at maturity were most affectedby a 10 d period of high temperature commencing 7–10 dafter anthesis However, if dry-matter accumulation between thestart of a treatment and grain maturity was used as a base forcomparison, the response was more uniform throughout development,with a peak in sensitivity 25 d after anthesis Although grainposition within an ear did not have a large effect on the responseto temperature, grains from the basal spikelets were more sensitivethan those from the apex, and the upper floret grains of a spikeletwere more sensitive to high temperature than those at the base There is a need to obtain, for a range of cultivars, more comprehensivedata on the effect of the timing and degree of temperature stressfollowing anthesis, for use in interpreting the response torising temperatures late in the development of the crop in thefield Triticum aestivum L, wheat, temperature, grain development     

The Effects of Floret Sterilization on Grain Number and Grain Weight in Wheat Ears

This paper describes the effects of sterilizing certain floretson the development of others within the ears of wheat, cultivarMaris Ranger. Sterilization of all the florets in spikelets2, 4, 6, and 8 (numbered from the base of the ear upwards) ledto more grain setting and greater grain growth in the untreatedspikelets. These compensatory increases were insufficient toprevent a depression in the yield of grain per ear. Sterilizationof more than one of the basal florets of spikelet 8 led to amore frequent setting of the grain in the distal florets onthat spikelet and to the centrally positioned grain becomingheavier. The physiological basis for the inhibitory influences of theolder and developing grain on the unfertilized florets and youngergrain is discussed.     

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.     

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     

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     

Distribution of carbon-14 assimilated by wheat awns

The pattern of distribution of carbon assimilated by awns was investigated in two lines of Triticum aestivum. Single awns on basal florets of spikelets in the central part of the ear were dosed with ¹⁴CO₂. Five days after dosing, 99% of the carbon-14 recovered was in the spikelet bearing the awn. Of the carbon-14 exported from the treated awn 57% went to the grain of the first floret, 1% to the second, 28% to the third and 7% to the fourth.     

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     

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.     

The effect of growth retardant application on floret site utilisation and assimilate distribution in ears of perennial ryegrass cv. S.24

Application of the growth retardant paclobutrazol (PP333), at 2 kg a.i. ha^-1 at spikelet initiation to plots of perennial ryegrass cv. S.24 in 1981 and 1982 significantly increased the number of seeds per spikelet present at final harvest by reducing the number of seeds aborted during seed development. Distribution of florets and seeds per spikelet was altered by PP333, as both basal and penultimate spikelets contained more florets and seeds than did those of untreated plants. Seed weight and germination were increased in florets of penultimate spikelets, although PP333 application delayed maturity by 3–5 days. In untreated plants, assimilate recovery was significantly lower from the terminal section of the ear, whereas in PP333 treated plants, no differences were found between basal, intermediate or terminal sections of the ear. PP333 increased assimilate demand at all sections of the ear when the ear and leaves were fed. The implications of this are discussed.     

Development and Floret Survival in Wheat Ears With Supernumerary Spikelets

In the supernumerary spikelet wheat, AUS159I0, the supernumeraryspikelet primordia appeared just after the ear reached the terminalspikelet stage. Appearance of the primordia of the multiplesessile spikelets preceded that of indeterminate rachilla spikelets.Supernumerary spikelets had a lower number of potentially fertileflorets per spikelet than normal (non-supernumerary) spikeletsin the ear and thus a smaller number of grains per spikelet.Mean weight per grain in the supernumerary spikelet wheat waslower than in the cultivar, Meering, without supernumerary spikelets.Total grain number in the supernumerary spikelet ear was greaterthan in the normal ear despite the lower spikelet fertilityin the former. Within the supernumerary spikelet ear the multiplesessile spikelets had a higher number of grains per spikeletand mean weight per grain than the indeterminate rachilla spikelets.It appears possible to improve the productivity of the supernumeraryspikelet ear by breeding for reduced expression of the indeterminaterachilla spikelets. Wheat, ear development, floret survival, supernumerary spikelets, grain number     

The Effect of Temperature on Seed Set and Seed Development in Detached Spikelets of Perennial Ryegrass (Lolium perenne L.)

In perennial ryegrass seed yield is low. Pollination, fertilizationand early stages of seed development are critical periods forthe realization of the yield potential. As intact plants aredifficult to handle, we used detached spikelets to study effectsof temperature and genotype on seed set and seed development. Plants of four cultivars were grown in pots. Just prior to flowering,spikelets were detached from the ears and put on water. Afterpollination, various spikelets of each plant were placed atdifferent temperatures (14/14, 17/12 or 20/15°C). Sevendays after pollination the percentage seed set and the lengthof the developing seeds (caryopses) were assessed. Only floretsin the four basal positions were used. For each cultivar an analysis of variance for unbalanced datawas carried out. There were highly significant difference amonggenotypes for seed set and caryopsis length. There was no evidencefor genotypic variation for tolerance to low temperatures. Neithertemperature nor floret position had an effect on seed set. Thecaryopsis length, however, increased with increasing temperatureand basal florets produced longer caryopses than distal florets.The detached spikelets proved to be a useful experimental tool.Copyright1993, 1999 Academic Press Lolium perenne L., perennial ryegrass, seed yield, seed set, detached spikelets culture, temperature, floret position     

Changes in Cell Number and Cell Division Activity during Endosperm Development in Allohexaploid Wheat, Triticum aestivum L.

Nuclear or cell number, and the mitotic index, were recordedin endosperms of Triticum aestivum cv. Mardler to test if aparticular stage of endosperm development was critical in determiningthe final grain weight. The basal four florets of emasculatedspikelets (controls), and the third and fourth florets of spikeletswhere the two basal ovaries were removed (ovary-removed), weresampled at various times up to 360 h after hand-pollination.The coenocytic phase in endosperms ended about 84 h after pollinationregardless of both grain position and the treatment. The onsetof the cellular stage was characterized by the final large fluctuationsin the mitotic index reflecting the culmination of the synchronousnuclear division of the coenocytic stage. Thereafter, the mitoticindex fluctuated with smaller amplitudes and, by 216 h afterpollination, was < 1%. Neither floret position in the spikeletnor the treatment affected the pattern of alteration to themitotic index. However, ovary removal from first and secondflorets resulted in significantly heavier grains and higherendosperm cell number in the 3rd and 4th florets compared withthe controls. In all florets, mean endosperm cell number peakedat 280 h but decreased by 360 h after pollination. At this time,the mean cell numbers in endosperms of the 3rd and 4th floretsof ovary-removed spikelets were significantly higher than inthe corresponding endosperms in the controls. Thus, a key contributoryfactor in determining the final endosperm cell number may bethe number of cells which are lost during the late period ofthe cellular stage of endosperm development. Key words: Endosperm cell number, florets, grain weight, mitotic index, Triticum aestivum     

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     

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.     

The Effect of Speckled Leaf Blotch on Apical Development and Yield in Winter Wheat in New Zealand

Speckled leaf blotch (Mycosphaerella graminicola) reduced yieldin a winter wheat trial by 18 per cent. Detailed analyses ofthe effects of disease on yield components showed that diseasereduced both grain number per ear and grain weight. The reductionin grain number per ear was due to a decrease in grain numberper spikelet, and this occurred for both main stem and tillers,and occurred equally at each spikelet position on the ear. The effect of disease on grain number was traced back to anearlier reduction in floret primordium number per ear, especiallyfloret number per spikelet. It was suggested that floret primordiumproduction was affected by the reduced assimilate supply tothe developing apex. The effects of disease on yield could therefore be attributedto effects on plant development both before and after anthesis.The disease effects on yield were not necessarily associatedwith the time of maximum disease severity. Plots with the early phase of disease epidemic only showed asignificant reduction in yield, but there was some evidenceof compensation for early reductions to yield potential in thelater-determined yield components. We suggest, therefore, thatan effective disease-control programme must take into accountthe possible early effects of disease on yield potential. Mycosphaerella graminicola, speckled leaf blotch, winter wheat, yield loss, apical development     

Effect of awns and drought on the supply of photosynthate and its distribution within wheat ears

The presence of awns doubled the net photosynthetic rate of wheat ears and also increased the proportion of ¹⁴CO₂ assimilated by the ear that moved to the grain. The effect of water supply on photosynthesis and movement of assimilates was greater for leaves than ears, so that drought increased the proportion of assimilate contributed by ear photosynthesis to grain filling from 13% to 24% in the awnless ears, and from 34% to 43% in the awned ears. ¹⁴C assimilated by the ears was most important to the economy of the upper spikelets and to the distal florets in each spikelet, whereas flag leaf assimilate went mainly to the spikelets in the lower half of the ear, and to the proximal florets. Awns increased grain yield in the dry but not in the irrigated treatment, despite the large contribution of awned ears to grain filling. Either the supply of assimilate did not limit grain yield when water supply was not limiting, or there were compensating disadvantages to awns. However, they did not seem to have any adverse effect on the development of the upper florets, nor did they reduce grain number per ear.     

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.     

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.     

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