Interrelationship between Meristematic Regions of Developing Inflorescences of Four Cereal Species

A cultivar from each of four cereal species (Avena sativa L.,cv. Swan, Hordeum vulgare, L., cv. Clipper, Triticum aestivumL., cv. Gabo, and Secale cereale L., cv. South Australian Rye)was grown in controlled environment chambers in a 10-h photoperiod(short days) or 10-h photoperiod supplemented with a 6-h extensionby incandescent light. The developmental morphology of the inflorescenceswas followed to ascertain whether there were any common developmentalinterrelationships between the species. Inflorescence internodeelongation was initiated when the floret initial first appeared,irrespective of whether it occurred on the most advanced lateralspikelet or on the terminal spikelet of the rachis. The glumes(infertile bracts) of the terminal spikelet of the rachis wereinitiated when the first second-order inflorescence branch appeared,irrespective of whether the second-order inflorescence branchwas a floret initial or a lateral spikelet, as in Triticum sp.,or an inflorescence (panicle) branch, as in Avena sp. Cessationof the activity of the apical meristem, as measured by primordiumformation, was not correlated with any particular stage of floraldevelopment but appeared to be due to a lack of nutrients causedby an increasing competitiveness for the available nutrientsfrom the developing spikelets which are situated closer to thevascular system than the apical meristem.     

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     

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     

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     

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     

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.)     

Culture of Barley Anthers in Conditioned Media

High yielding anther cultures of Hordeum vulgare cv. Sabarlisare obtained at inoculation densities of 10—20 per mlby use of media previously conditioned by Sabarlis anthers.To achieve these high yields anthers at the mid-unicellularpollen stage (stage 2), stressed in the excised spike for 14d at 7 °C, are necessary, but for conditioning of mediaolder anthers may be used with or without the stress pretreatment.Conditioning for 7 d by anthers at young bicellular pollen stages(stages 5—6) is highly effective. Hormones supplied inthe medium interact synergistically with the conditioning factor. Sabarlis ovaries are shown to be even more effective for conditioningthan anthers, whereas glumes and other parts of the spike arerelatively ineffective. Anthers of oats, rye, wheat, maize,tobacco, and one other genotype of barley are also less effectivethan Sabarlis anthers. The improved method of anther culture is more efficient thanspike culture for the production of pollen callus in Sabarlisbarley.     

Pollination Biology in a TropicalAcaciaHybrid (A. mangiumWilld. xA. auriculiformisA. Cunn. ex Benth.)

Floral biology and pollination mechanism of theAcaciahybrid(A. mangiumWilld. xA. auriculiformisA. Cunn. ex Benth.) growingin Thailand are investigated using light and electron microscopy.The hybrid is andromonoecious. A floral spike consists of about150 loosely arranged flowers. Flowers are cream coloured, fragrantand have no floral nectaries. The pistil has a solid style witha smooth, wet stigma and amphitropous ovules with immature integuments.The anther consists of eight loculi, each bearing only one 16-grainpolyad. The flowers are weakly protogynous. Anthesis is completeat 0500–0600 h but peak female receptivity begins at 0200–0300h and is completed that day. The stigmatic exudate is of thelipophilic type and is secreted from the stigmatic cells bya holocrine mechanism. Pollen is the main floral reward forthe insect pollinators. There are several floral characteristicswhich facilitate pollen transfer from anthers and depositionon stigmas.Apis melliferaandCeratinasp. are the most effectivepollinators because they are the most common visitors and carrya heavy load of hybrid polyads. However, their behaviour inforaging for pollen in the same tree and weak protogynous dichogamymay promote self-pollination in the hybrid. The hybrid has lowpollination success due to low pollinator number. An increasein exposure time of flowers to pollinators or pollinator numbermay increase pollination success but may not affect the rateof pollen deposition on stigmas due to the relatively smallsize of the stigma in relation to the polyad.Copyright 1998Annals of Botany Company Acaciahybrid,Acacia mangium, Acacia auriculiformis, pollination, pistil receptivity, anthesis, pollinators.     

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.     

Use of Longitudinally Bisected Pistils of Lilium longiflorum for Studies on Self-Incompatibility

The longitudinally bisected pistil of Lilium longiflorum cv.Hinomoto was shown to be useful for the study of self-incompatibility.When pollen grains of cv. Hinomoto or cv. Georgia were placedon the stigma of the bisected pistil, pollen tube elongationof each cultivar occurred with almost the same time course aswhen placed on the stigmaof the whole pistil. Tube elongation of cv. Hinomoto was retarded after 24 hours(self combination), whereas the tubes of cv. Georgia elongatedwell (cross combination). The selfincompatibility reaction detectedas retardation of pollen tube elongation occurred in all portionsof the inner surface of the stylar canal; it was not restrictedto a specific portion of the style. ¹Present address: Laboratory of Pomology, Faculty of Horticulture,Chiba University, Matsudo, Chiba 271, Japan. (Received October 20, 1982; Accepted February 9, 1983)     

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.     

Quantifying relationships between rooting traits and water uptake under drought in Mediterranean barley and durum wheat

In Mediterranean regions drought is the major factor limiting spring barley and durum wheat grain yields. This study aimed to compare spring barley and durum wheat root and shoot responses to drought and quantify relationships between root traits and water uptake under terminal drought.One spring barley（Hordeum vulgare L. cv. Rum） and two durum wheat Mediterranean cultivars（Triticum turgidum L. var durum cvs Hourani and Karim） were examined in soil‐column experiments under well watered and drought conditions. Root system architecture traits, water uptake, and plant growth were measured. Barley aerial biomass and grain yields were higher than for durum wheat cultivars in well watered conditions. Drought decreased grain yield more for barley（47%） than durum wheat（30%, Hourani）. Root‐to‐shoot dry matter ratio increased for durum wheat under drought but not for barley, and root weight increased for wheat in response todrought but decreased for barley. The critical root length density（RLD） and root volume density（RVD） for 90% available water capture for wheat were similar to（cv. Hourani） or lower than（cv. Karim） for barley depending on wheat cultivar. For both species, RVD accounted for a slightly higher proportion of phenotypic variation in water uptake under drought than RLD.     

Stereological Analysis of Floral Development and Quantitative Histochemistry of Nucleic Acids in Fertile and Base-sterile Varieties of Wheat

The speltoid series of mutations provides a genetic tool forinvestigating the control of flower development. In fertileGabo wheat and in St₂ and St₃ basal sterile speltoids, the changesin RNA staining patterns have been followed histochemicallyas a marker of cellular activity. Sterile floret sites are characterizedby a reduced RNA content. Use of dual-wavelength microspectrophotometryhas provided evidence that RNA content is similar in lemma primordiasubtending presumptive fertile and sterile primordia, but isstrikingly different in the floral meristems in the two types.The total nucleic acid content of the floral primordia has beenrelated to the number of epidermal cells in the subtending lemmaprimordium as a marker of development, using a linear modelanalysis of covariance. There is a linear increase in nucleicacid content during fertile floret development, but no significantincrease in RNA content in sterile St₃ primordia. The rate constantis only 1/10th to 1/20th of that of Gabo. A stereological analysisshows that cell number in the floral meristems of Gabo increasesexponentially during development. In contrast, in St₃, whilethere is a significant increase in cell number, it is at a drasticallyreduced rate. The intercept values are close to zero, indicatingthat only one or two cells initiate floral meristem development.The fertility-controlling alleles exert their effect prior tothe appearance of a visible floral primordium, and probablyafter initiation of the lemma. Triticum aestivum, wheat, floral development, histochemistry, nucleic acids, speltoids     

Stigma-surface Esterase Activity and Stigma Receptivity in Some Taxa Characterized by Wet Stigmas

Stigma-surface esterase activity and stigma receptivity througha sequence of developmental stages of the pistil have been studiedin four taxa characterized by having wet stigmas — Petuniahybrida, Nicotiana tabacum, Crinum defixum and Amaryllis vittata.The style is solid in the first two and hollow in the lattertwo taxa. In all the taxa, stigma—surface esterase couldbe detected in a thin surface layer (pellicle) from a very earlystage of pistil development, irrespective of the presence orabsence of the exudate. However, the taxa showed variation instigma receptivity. In Petunia and Nicotiana, stigmas from pistilsof all the stages supported pollen germination and tube growth.In Amaryllis and Crinum, stigmas of only the mature pistils,when the exudate is present on the stigma, supported normalpollen germination and tube growth. It is inferred that in taxacharacterized by a wet stigma and solid style, the factors requiredfor pollen germination are present from an early stage of pistildevelopment and the exudate per se is not involved in pollengermination. In taxa characterized by a wet stigma and hollowstyle, however, the pellicle does not carry the factors requiredfor pollen germination and tube growth; they appear to be presentin the exudate. Petunia hybrida Vilm, Nicotiana tabacum L., Crinum defixum, Ker-Gawl, Amaryllis vittata Ait., tobacco, pollination, pollen germination, stigmatic exudate, stigma receptivity, stigma-surface esterase, esterase activity     

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.     

Comparative mapping of HKT genes in wheat, barley, and rice, key determinants of Na+ transport, and salt tolerance

Salt tolerance of plants depends on HKT transporters (High-affinityK⁺ Transporter), which mediate Na⁺-specific transport or Na⁺-K⁺co-transport. Gene sequences closely related to rice HKT geneswere isolated from hexaploid bread wheat (Triticum aestivum)or barley (Hordeum vulgare) for genomic DNA southern hybridizationanalysis. HKT gene sequences were mapped on chromosomal armsof wheat and barley using wheat chromosome substitution linesand barley–wheat chromosome addition lines. In addition,HKT gene members in the wild diploid wheat ancestors, T. monococcum(A^m genome), T. urartu (A^u genome), and Ae. tauschii (D^t genome)were investigated. Variation in copy number for individual HKTgene members was observed between the barley, wheat, and ricegenomes, and between the different wheat genomes. HKT2;1/2-like,HKT2;3/4-like, HKT1;1/2-like, HKT1;3-like, HKT1;4-like, andHKT1;5-like genes were mapped to the wheat–barley chromosomegroups 7, 7, 2, 6, 2, and 4, respectively. Chromosomal regionscontaining HKT genes were syntenic between wheat and rice exceptfor the chromosome regions containing the HKT1;5-like gene.Potential roles of HKT genes in Na⁺ transport in rice, wheat,and barley are discussed. Determination of the chromosome locationsof HKT genes provides a framework for future physiological andgenetic studies investigating the relationships between HKTgenes and salt tolerance in wheat and barley. Key words: Barley, comparative mapping, HKT, rice, salt tolerance, sodium transport, wheat     

The Control of Flowering in Wheat and Barley: What Recent Advances in Molecular Genetics Can Reveal

The combined forces of developmental biologists, studying primordiuminitiation at the stem apex, and mathematical modellers, developingsimulations of crop growth and development, have brought aboutconsiderable advances in the understanding of the control offlowering in wheat and barley. Nevertheless, there are stillmajor gaps in this understanding including: what determinesthe basic rate of development (magnitude of the phyllochronor plastochron); how temperature and photoperiod interact tobring about the transition from vegetative to reproductive development;and how flowering occurs eventually in the absence of inductiveconditions. Although geneticists have tended to measure cerealflowering in terms of ‘days from sowing or emergence toheading’, results of studies using aneuploids and molecularmarkers are compatible with the roles for photoperiod and low-temperaturevernalization established in purely-physiological or developmentalinvestigations. They have also revealed the existence of ‘earlinessperse’loci, whose detailed roles have yet to be established.Progress towards isolating and characterizing wheat and barleyloci is hampered by the poor resolution of mapping (locationto a precision of tens of thousands of base pairs). Neitherof these broad approaches promises a rapid resolution of thefactors controlling the induction of flowering. Two expandingareas of molecular genetics now provide potential for greaterunderstanding of cereal flowering. First, the extensive homoeologyamong members of the Gramineae can be employed to establishthe existence and location of genes or quantitative trait lociin rice which correspond to controlling loci in wheat or barley.Since the rice genome is 1/30th of the size of the wheat genome,the accuracy of mapping loci can be much higher, and there isgreater potential for precise location of loci using techniquessuch as chromosome walking. With the ultimate cloning of individualgenes, and the isolation of gene products, the relative rolesof the 20 loci apparently involved in the induction of floweringof wheat could be explored. However, progress in the moleculargenetics ofArabidopsis(the second area) may provide a more rapidroute to understanding the control of flowering in cereals forseveral reasons: its small genome (1/4 that of rice); the likelihoodof extensive homoeology with cereals, in spite of differencesin codon usage between monocots and dicots; the existence ofa wide range of flowering-time mutants; and the control of floralinduction by a similar range of environmental factors includingphotoperiod and low temperature. It is likely that the MCDK(Martinez-Zapater, Coupland, Dean and Koornneef, 1994. In: MeyerowitzEM, Somerville CR.Arabidopsis.New York: Cold Spring Harbor Laboratory,403–433) model, formulated to explain the genetic andenvironmental control of flowering inArabidopsis,could be employedusefully in the formulation of experimental work on floweringin wheat and barley. This paper reviews these issues, payingparticular attention to the significance of ‘earlinessperse’ loci and the ‘constitutive floral pathway’for wheat and barley.Copyright 1998 Annals of Botany Company Wheat, barley, rice,Arabidopsis,flowering, photoperiod, vernalization, genetics, development.     

Male Sterility and Anther Wall Structure in Copper-deficient Plants

Anther development and pollen sterility were followed in plantsof wheat, oat, barley, sweetcorn, sunflower, petunia and subterraneumclover grown at a range of copper supplies. Copper-deficientplants had increased pollen sterility. Lignified wall thickenings were reduced or absent in the endotheciaof anthers from Cu-deficient plants. Reduced seed set may resultboth from reduced pollen fertility or failure of the stomiato rupture due to decreased lignification of anther walls. Triticum aestivum L., wheat, Hordeum vulgare L., barley, Avena sativa L., oat, Zea mays L., corn, sweetcorn, maize, Helianthus annuus L., sunflower, Petunia hybrida L., Trifolium subterraneum L., subterranean clover, male sterility, anther development, copper deficiency     

The Influence of Brief Exposures to Short Days on the Growth and Development of Barley

Plants of barley (Hordeum vulgare L. cultivar Proctor) otherwisegrown, in long days (16 h) were given a 2-week exposure to shortdays from the following stages of main shoot development: (A)double ridge, (B) stamen initial, (C) awn initial, and (D) alater stage of awn differentiation. Treatment B led to severeinhibition and ultimate death of the main shoot ear and treatmentsC and D to the almost complete failure of certain ears to setgrain. In main shoot ears the reduction in floret fertilitywas shown to be due to male sterility. The effects on shootmortality and floret fertility are new and have not been observedby other workers when plants were grown in continuous shortdays. They therefore appear to depend on an exposure to longdays before the treatment with short days. In contrast, theeffects of the short days on vegetative characters, such asthe number, length, and width of leaves on the main shoot andthe numbers of tillers produced, were similar to those whichhave been observed under continuous short days.