Photoperiod and Vernalization Responses in Triticum turgidumxT. tauschii Synthetic Hexaploid Wheats

Studies of synthetic hexaploid wheat developed from Triticumturgidum(AABB genomes) and T. tauschii(DD genome) can provideinformation on potentially useful characters in T. tauschiiand/or T. turgidum for genetic improvement of hexaploid wheat(T. aestivum). Synthetic hexaploid wheats and the T. turgidumand T. tauschii parents were assessed for their developmentalresponses to photoperiod and vernalization for days to ear emergence,final leaf number and the number of spikelets per spike. Theresponses to photoperiod and vernalization of the synthetichexaploids were generally intermediate between those of theparents but in some instances the levels of expression exhibitedby the T. tauschii or T. turgidum parents were epistatic inthe synthetic hexaploids. The relatively strong photoperiodresponse of the T. tauschii accessions was not expressed inthe synthetic hexaploids, but rather the synthetic hexaploidsreflected the photoperiod response of the respective T. turgidumparents. The synthetic hexaploids had vernalization responsesstronger than those of the T. turgidum and bread wheats usedin the study. The expression of ear emergence in response tovernalization of these synthetic hexaploids appeared to be modifiedby the T. turgidum parent. Copyright 2001 Annals of Botany Company Photoperiod, synthetic hexaploids, Triticum aestivum, Triticum tauschii, Triticum turgidum, vernalization     

Expression of Vernalization Requirement and Spikelet Number in Synthetic Hexaploid Wheats and their Triticum tauschii and Tetraploid Wheat Parents

Vernalization requirement, as measured by days from sowing toear emergence (plants grown under an 18-h photoperiod), andspikelet number per ear were recorded for 17 synthetic hexaploidwheats and the six tetraploid (Triticum durum) and the ninediploid T. tauschii parents used to synthesize them. The tetraploid parents and the synthetic hexaploids had springphenotypes (little or no vernalization requirement) whereasthe T. tauschii parents were all winter types (strong vernalizationrequirement). The tetraploid wheats and the synthetic hexaploidsreached ear emergence 50·3 to 63·8 d and 58·2to 75·3 d after sowing, respectively, while the T. tauschiilines reached ear emergence 114·3 to 179·5 d aftersowing. The spring habit of the synthetic hexaploids demonstrates theepistasis of spring over winter habit. It is considered thatwith a presumed single vrn locus in the diploid species T. tauschiithe range of ear emergence in these lines is consistent withthe action of multiple alleles at that locus. Although there was no general epistasis for spikelet number,the tetraploid parents appear to be exerting more influenceover spikelet number in the synthetic hexaploids than T. tauschii.The well established association between the duration from sowingto ear emergence and spikelet number was not evident eitherwithin each ploidy group or when the 32 lines were consideredtogether. Triticum tauschii, Triticum durum, hexaploid wheat, spikelet number, vernalization requirement     

The Growth and Development of the Wheat Apex: The Effects of Photoperiod on Spikelet Production and Sucrose Concentration in the Apex

Spring wheat (Triticum aestivum cv. Warimba) plants were grownin a controlled environment (20°C) in two photoperiods (8or 16 h). In the first instance, plants were maintained in eachof the photoperiods from germination onwards at the same irradiance(375 µE m^–2 s^–1). In the second case, allplants were grown in a long photoperiod until 4 days after double-ridgeinitiation when half the plants were transferred to a shortphotoperiod with double the irradiance (16 h photoperiod at225 or 8 h at 475 µE ^–2 s^–1). The rates of growth and development of the apices were promotedby the longer photoperiod in both experiments. Shoot dry weightgain was proportional to the total light energy received perday whereas the dry weight of the shoot apex increased withincreasing photoperiod even when the total daily irradiancewas constant. The principal soluble carbohydrate present in the shoot apexwas sucrose, although low concentrations of glucose and fructosewere found in the apices of long photoperiod plants late indevelopment. Sucrose concentration was invariably greater inthe slow-growing apices of short photoperiod plants, but roseto approach this level in the long photoperiod plants when theterminal spikelet had been initiated. Triticum aestivum, wheat, apex, spikelet initiation, photoperiod, flower initiation     

Influence of Photoperiod on Culm Elongation and Apical Development in Semi-dwarf and Standard-height Wheats

Flower initiation (FI) coincided with the commencement of culmelongation under both long (18 h) and normal (104–144h) photoperiod in eight spring wheats, including both gibberellicacid-sensitive and -insensitive types, which differed widelyin photoperiod sensitivity At FI the apex was significantly (P = 005) higher (above ground)in three of the wheats under long, compared with normal photoperiod;with no difference between the remaining five. Differences inresponse were not related to photoperiod response or gibberellicacid-sensitivity/insensitivity differences between the wheats. Long photoperiod prolonged the phase from terminal spikeletinitiation (TSI) to anthesis (A) in all the wheats, except Sunset(with the greatest photoperiod insensitivity), with no cleardifferences in response between semi-dwarf and standard-heightwheats. Respective rates of culm elongation from FI to TSI were lowerunder normal, compared with long, photoperiod in all varieties.That from TSI to A was unaffected by photoperiod, except inSunset when it was significantly (P = 001) slower under long,compared with normal photoperiod. Rate of culm elongation from FI to TSI across cultivars andphotoperiods was inversely related to spikelet number per head(r = –053, P = 005) but not to rate of spikelet initiation(r = –014 n.s.). Gibberellin-sensitivity, spikelet number, flower initiation, terminal spikelet initiation     

Expression of Vernalization Genes in Near-isogenic Wheat Lines: Effects of Night Temperature

Four near-isogenic lines of wheat (Triticum aestivum L.em Thell)were used to compare selected night temperatures for their effectivenessas vernalizing temperatures. All treatments (conducted withina phytotron) had a common day temperature of 20 °C for 12h and night temperatures were 4, 7, 10, 13 and 20 °C. Interpretationof results for reproductive development was confounded by threeinteracting factors, their relative importance varying withgenotype. Firstly, development rate was generally slower atlower night temperatures. Secondly, in contrast, there was atendency for lower night temperatures to hasten developmentrate if vernalization requirements were satisfied. Thirdly,the lower night temperatures provided a more favourable environmentfor leaf production such that for some genotypes, vernalizedplants had higher final leaf numbers than unvernalized plants.Only for the genotype with the strongest vernalization response(vrn1 vrn2) did hastening of development due to vernalizationoverride any delaying effects. For this genotype, 4, 7 and 10°C were vernalizing temperatures. For the other three genotypes,any hastening of development due to vernalization was outweighedby delaying effects of lower night temperatures. Spikelet numberand days to anthesis were positively correlated in three ofthe four genotypes. It appeared that differences in spikeletnumber were a direct result of night temperature influencingthe duration of the spikelet phase and/or rate of spikelet initiation.Plant size at flowering was determined by the differential effectsof night temperature on growth and development rates. Triticum aestivum L., wheat, vernalization, night temperature, isogenic lines     

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     

Developmental Regulation of Low-temperature Tolerance in Winter Wheat

Vernalization and photoperiod genes have wide-ranging effectson the timing of gene expression in plants. The objectives ofthis study were to (1) determine if expression of low-temperature(LT) tolerance genes is developmentally regulated and (2) establishthe interrelationships among the developmental stages and LTtolerance gene expression. LT response curves were determinedfor three photoperiod-sensitive LT tolerant winter wheat (Triticumaestivum L. em Thell) genotypes acclimated at 4 °C under8 h short-day (SD) and 20 h long-day (LD) photoperiods from0 to 112 d. Also, three de-acclimation and re-acclimation cycleswere used that bridged the vegetative/reproductive transitionpoint for each LD and SD photoperiod treatment. A vernalizationperiod of 49 d at 4 °C was sufficient for all genotypesto reach vernalization saturation as measured by minimum finalleaf number (FLN) and confirmed by examination of shoot apicesdissected from crowns that had been de-acclimated at 20 °CLD. Before the vegetative/reproductive transition, both theLD- and SD-treated plants were able to re-acclimate to similarLT₅₀(temperature at which 50% of the plants are killed by LTstress) levels following de-acclimation at 20 °C. De-acclimationof LD plants after vernalization saturation resulted in rapidprogression to the reproductive phase and limited ability tore-acclimate. The comparative development of the SD (non-flowering-inductivephotoperiod) de-acclimated plants was greatly delayed relativeto LD plants, and this delay in development was reflected inthe ability of SD plants to re-acclimate to a lower temperature.These observations confirm the hypothesis that the point oftransition to the reproductive stage is pivotal in the expressionof LT tolerance genes, and the level and duration of LT acclimationare related to the stage of phenological development as regulatedby vernalization and photoperiod requirements. Copyright 2001Annals of Botany Company Triticum aestivum L., wheat, low-temperature tolerance, vernalization, photoperiod, phenological development     

Expression of Vernalization Genes in Near-lsogenic Wheat Lines: Duration of Vernalization Period

Vernalization periods ranging from 0 to 11 weeks at 4 °Cwere used to study the reproductive development of four near-isogeniclines of wheat (Triticum aestivum L. em. Thell). From the resultsfor days to anthesis two types of gene action were identified,a threshold (all-or-nothing) response (vrn3 and/or vrn4) anda cumulative (graded) response (vrn1). The action of anothergene (vrn2) intensified these two responses. Based on the actionof genes, a model relating days to anthesis to genotype wasderived. Final leaf number and days to anthesis were shown tobe closely related after adjusting for differences due to theduration of vernalization treatment. No relationship betweendays to anthesis and spikelet number was observed. This studyemphasises the need to understand vernalization at the levelof the gene in terms of responses and interactions. Such knowledgeshould enable the plant breeder to predict and more preciselycontrol reproductive development. Triticum aestivum L., wheat, vernalization, gene action, isogenic lines     

Development in wheat as affected by timing and length of exposure to long photoperiod

Seeds of a spring wheat (Triticum aestivum L. cv. ‘Condor’)were vernalized and then grown at 19C in two naturally–litenvironments, one with a moderate (12 h) and the other withlong (18 h) photoperiod. Treatments consisted of transfers ofplants from the moderate to the long photoperiod chamber ondifferent occasions, or for periods of different durations.The main objectives were to determine whether wheat developmentresponds to current and previous photoperiodic environmentsand whether there is a juvenile phase when the plants are insensitiveto photoperiod. Plants under constant 18 h photoperiod had fewerleaves which appeared faster than those under constant 12 hphotoperiod (i.e. phyllochron was increased from 4.4 to 5.1d leaf^–1). Plants transferred from 12 h to 18 h photoperiodat terminal spikelet appearance (TSA) reached anthesis 4 d earlierthan plants retained at 12 h, while plants under continuouslong photoperiod (18 h) completed this phase most rapidly. Thus,there was some evidence for a historic effect of photoperiodon development. Exposure to long photoperiod during the first 5 d after plantemergence accelerated the rate of development towards anthesis,suggesting that there was no juvenile period of photoperiodicinsensitivity. There were, however, changes during ontogenyin the degree of sensitivity to long photoperiod, increasingfrom seedling emergence to a maximum c. 15 d later, and thendecreasing again. Although all treatments were imposed beforeTSA, the response was not limited to the pre-TSA phase, suggestingthat well before the terminal spikelet appeared, the plant wasalready committed to the initiation of this spikelet. Spikeletnumber decreased with delayed transfer to long photoperiod witha minimum for plants transferred to long days from 16-20 d afterseedling emergence. Additionally, there was a trend for an increasein the rate of leaf appearance (decrease in phyllochron) whenthe plants were exposed to long days between 10 and 35 d afterseedling emergence. Although the differences were small, whenconsidered in conjunction with the effects on final leaf numberthey become important in explaining differences in time to anthesis. Key words: Development, flowering, leaf number, photoperiod, phyllochron, Triticum aestivum     

Duration and Rate of Development Phases in Wheat in Two Environments

Six cultivars of bread wheat (Triticum aestivum ssp. vulgareL. amend. Thell.) of diverse climatic origin and different developmentalpatterns were studied under two environments, (a growth roomand an outdoors sowing) for the duration and rate of completionof their developmental phases. The need for vernalization in the cultivar Cappelle Desprezsubstantially increased the length of the vegetative phase,particularly in the growth room. Large differences in the durationof reproductive initiation and stem elongation phases betweenCappelle Desprez and the other cultivars in the growth roomsowing suggests an influence of veralization beyond the vegetativephase. Differences between the two environments influenced the durationof all phases of development, giving pronounced between-cultivarvariation in both the stem elongation and ripening phases. Rates of reproductive initiation and stem elongation for thecultivars, within and between the two environments appearedto be largely independent. The rate of spikelet initiation wassignificantly decreased in the growth room compared with theoutdoor sowing. The duration of the phases of development withineach cultivar appeared to be independent of each other, indicatingthe possibility for adjusting the rate, or duration, of a phaseof development comparatively free of a compensatory change inthe rate, or duration, of other phases. Triticum aestivum ssp, vulgare, wheat, vegetative phase, flower initiation, vernalization, photoperiodism     

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 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     

Effects of N and P nutrition on spike development in spring wheat

The effects of N and P on the number of spikelets of spring wheat (Triticum aestivum L.), grown in nutrient solution, were studied under 8 h and 16 h photoperiods. The effect of P was apparent only at a high rate of N supply and the effects of N were increased significantly at a high rate of P supply. Increasing N supply increased the number of spikelets due to a promotion of the rate of spikelet initiation. It also increased the leaf-blade area and the dry matter weight of the plants at the stage of terminal spikelet initiation. These effects of N were much greater under the short photoperiod than under the long photoperiod. The practical significance of these findings for winter-grown wheat in temperate regions is pointed out.     

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     

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.)     

Development Rate in Wheat as Affected by Duration and Rate of Change of Photoperiod

A field study was conducted to test the hypothesis that wheatdevelopment rate responds to the rate of change of photoperiod.Two wheat cultivars (Condor and Thatcher) were sown on 18 Aug.1992 at Melbourne (38° S). Photoperiod was extended artificiallyto give five treatments up to terminal spikelet initiation (TS)viz.: natural photoperiod (rate of change of photoperiod, 2·3mind d^-1), two faster rates of change (9·8 and 13·1min d^-1) and two constant photoperiods of 14·0 and 15·5h. After TS, the two constant photoperiods were extended to15·0 and 16·5 h, respectively and treatments wererandomly re-allocated, i.e. some plots received different photoperiodregimes before and after TS. There were no significant differences among treatments in thelength of the period from sowing (S) to seedling emergence (E)phase, ranging from 15 to 16·3 d. The rate of developmentfrom E to TS responded to increases in photoperiod in both cultivars,increasing with average photoperiod across all treatments butthere was no effect of rate of change of photoperiod independentof its average photoperiod. The rate of development from TSto anthesis (A) did not show any trend with average photoperiod.This lack of effect of photoperiod on the period from TS toA contrasts with other results from the literature and possiblereasons for this conflicting result are discussed. Rate of changeof photoperiod did not affect the duration of the phase fromTS to A either. Therefore, the effect of photoperiod on theduration of the S-A period was strongly and positively correlatedto that of the length of the E-TS phase.Copyright 1994, 1999Academic Press Triticum aestivum L., wheat, phasic development, photoperiod, rate of change     

Temperature Response of Vernalization in Wheat: A Developmental Analysis

The vernalization response of wheat ( Triticum aestivum L.)was reinterpreted from a developmental perspective, using currentconcepts of the developmental regulation of wheat morphologyand phenology. At temperatures above 0 °C, the effects ofthe process of vernalization per se in wheat are confoundedby the effects of concurrent vegetative development. These effectsare manifested by differences in the number of leaves initiatedby the shoot apex prior to floral initiation, which in turnaffects the subsequent rate of development to ear emergenceand anthesis. Leaf primordia development during vernalizationand total leaf number at flowering were used to develop criteriato define both the progress and the point of saturation of thevernalization response. These criteria were then used to reinterpretthe results of Chujo ( Proceedings of the Crop Science Societyof Japan 35 : 177–186, 1966), and derive the temperatureresponse of vernalization per se for plants grown under saturatinglong day conditions. The rate of vernalization increased linearlywith temperature between 1 and 11 °C, such that the timetaken to saturate the vernalization response decreased from70 d at 1 °C to 40 d at 11 °C. The rate declined againat temperatures above 11 °C, and 18 °C was apparentlyineffective for vernalization. Total leaf number at saturation,however, increased consistently with temperature, as a resultof the balance between the concurrent processes of leaf primordiuminitiation and vernalization. Total leaf number at saturationincreased from 6 at 1 °C to 13.3 at 15 °C, which inturn influenced the time taken to reach ear emergence. The advantagesof using this developmental interpretation of vernalizationas the basis for a mechanistic model of the vernalization responsein wheat are discussed. Triticum aestivum L.; wheat; vernalization; rate; temperature; primordia; leaf number; flowering     

Differences in Zinc Efficiency among and within Diploid, Tetraploid and Hexaploid Wheats

Greenhouse experiments were carried out with six diploid, ninetetraploid and seven hexaploid wheats, including wild and primitivegenotypes, to study the influence of varied zinc (Zn) supplyon the severity of Zn deficiency symptoms, shoot dry matterproduction and shoot Zn concentrations. In addition to wildand primitive genotypes, one modern tetraploid cultivar withhigh sensitivity to Zn deficiency and two modern hexaploid cultivars,one highly sensitive to and one resistant to Zn deficiency,were included for comparison. Plants were grown for 44 d ina severely Zn-deficient calcareous soil, with (+Zn; 5 mg Znkg^-1soil) and without (-Zn) Zn fertilization. Visible Zn deficiencysymptoms, including whitish-brown necrotic patches on leaf blades,appeared very rapidly and severely in all tetraploid wheat genotypes.Compared with tetraploid wheats, diploid and hexaploid wheatswere less sensitive to Zn deficiency. With additional Zn, shootdry matter production was higher in tetraploid than diploidand hexaploid wheats. However, under Zn-deficient conditionstetraploid wheats had the lowest shoot dry matter production,indicating the very high sensitivity of tetraploid wheats toZn deficiency. Consequently, Zn efficiency expressed as theratio of shoot dry matter produced under Zn deficiency to Znfertilization, was much lower in tetraploid wheats than in diploidand hexaploid wheats. On average, Zn efficiency ratios were36% for tetraploid, 60% for diploid and 64% for hexaploid wheats.Differences in Zn efficiency among and within diploid, tetraploidand hexaploid wheats were positively related to the amount ofZn per shoot of the genotypes, but not to the amount of Zn perunit dry weight of shoots or seeds used in the experiments.The seeds of the accessions of tetraploid wild wheats containedup to 120 mg Zn kg^-1, but the resulting plants showed very highsensitivity to Zn deficiency. By contrast, hexaploid wheatsand primitive diploid wheats with much lower Zn concentrationsin seeds had higher Zn efficiencies. It is suggested that notonly enhanced Zn uptake capacity but also enhanced internalZn utilization capacity of genotypes play important roles indifferential expression of Zn efficiency. The results of thisstudy also suggest the importance of the A and D genomes asthe possible source of genes determining Zn efficiency in wheat.Copyright 1999 Annals of Botany Company Seeds, Triticum aestivum, Triticum monococcum, Triticum turgidum, zinc concentrations, zinc deficiency, zinc efficiency.     

Simulating the influence of vernalization, photoperiod and optimum temperature on wheat developmental rates

Background and Aims

Accurately representing development is essential for applying crop simulations to investigate the effects of climate, genotypes or crop management. Development in wheat (Triticum aestivum, T. durum) is primarily driven by temperature, but affected by vernalization and photoperiod, and is often simulated by reducing thermal-time accumulation using vernalization or photoperiod factors or limiting accumulation when a lower optimum temperature (T_optl) is exceeded. In this study T_optl and methods for representing effects of vernalization and photoperiod on anthesis were examined using a range of planting dates and genotypes.

Methods

An examination was made of T_optl values of 15, 20, 25 and 50 °C, and either the most limiting or the multiplicative value of the vernalization and photoperiod development rate factors for simulating anthesis. Field data were from replicated trials at Ludhiana, Punjab, India with July through to December planting dates and seven cultivars varying in vernalization response.

Key Results

Simulations of anthesis were similar for T_optl values of 20, 25 and 50 °C, but a T_optl of 15 °C resulted in a consistent bias towards predicting anthesis late for early planting dates. Results for T_optl above 15 °C may have occurred because mean temperatures rarely exceeded 20 °C before anthesis for many planting dates. For cultivars having a strong vernalization response, anthesis was more accurately simulated when vernalization and photoperiod factors were multiplied rather than using the most limiting of the two factors.

Conclusions

Setting T_optl to a high value (30 °C) and multiplying the vernalization and photoperiod factors resulted in accurately simulating anthesis for a wide range of planting dates and genotypes. However, for environments where average temperatures exceed 20 °C for much of the pre-anthesis period, a lower T_optl (23 °C) might be appropriate. These results highlight the value of testing a model over a wide range of environments.Key words: Wheat, Triticum aestivum, T. durum, air temperature, thermal time, shoot apex, phenology, growth stages, anthesis, flowering     

A Comparison of the Development of Multiflorous and Standard Wheats

This study was conducted to compare patterns of phenologicaldevelopment and agronomic traits of multiflorous and standardwheats (Triticum aestivum L. em. Thell.) when grown in environment-controlledgrowth rooms (20/15 °C, 16/8 h, day/night, 600 µmolm ² S ¹). Six winter and six spring habit genotypes were studiedin separate experiments. The winter multiflorous genotypes wereearlier than the winter standard genotypes. Their earlinessresulted in lower total biomass and grain yields, lower leafand tiller numbers per primary culm, and spikelet numbers perspike. Within the spring group, few differences were associatedwith spike type although the multiflorous genotypes had reducedgrowth as indicated by lower tiller and spike numbers per plant.Leaf emergence in the multiflorous genotypes, as in the standardgenotypes, was found to be a linear function of time. The tilleringpatterns of the multiflorous types were less pronounced thanfor the standards with lower maximum tiller numbers, and restricteddie-back and reinitiation. Floret numbers per spikelet werehigher in the multiflorous types due to higher initiation rates,longer initiation periods and/or reduced periods of die-back.The multiflorous trait was negatively associated with spikenumber and greater numbers of florets per spikelet did not alwaystranslate into greater numbers of kernels per spikelet. Becauseyield compensation was incomplete, yields per plant were lowerin the multiflorous genotypes than the standards. Triticum aestivum L. em, Thell., wheat, multiflorous trait, yield components, development