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.     

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     

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.     

Osmotic Adjustment in the Spikelets and Leaves of Wheat

The response of osmotic potential to changes in water potentialin the spikelets and leaves of wheat genotypes was studied ina controlled environment. While there was evidence of osmoticadjustment in the leaves of only some genotypes, it seemed tobe present in the spikelets of all genotypes. Spikelets alsodiffered from leaves in that osmotic potentials at full turgorwere usually higher. During grain growth, levels of osmoticpotential were observed in the spikelets of some genotypes whichwere up to 1·1 MPa greater than the corresponding waterpotential. Osmotic responses in both spikelets and leaves arediscussed in terms of responses expected from concentrationeffects only.     

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     

Changes in Levels of Endogenous Plant Hormones During Floret Development in Wheat Genotypes of Different Spike Sizes

The levels of endogenous plant hormones regulate floret development and degeneration, and thus grain set in flower crops. This study was undertaken to characterize the changes of endogenous hormone levels during floret development in three wheat (Triticum aestivum L.) genotypes: “97J1" with the highest grain set and fertile florets per spike, “H8679" with the lowest grain set and fertile florets per spike, and a medium, “YM158". The results showed that the peak level of ABA appeared between stamen and pistil differentiation and antherlobe formation of floret development, and the timing delayed with the size of spike (earliest in “H8679” and latest in “97J1”). From antherlobe formation to meiosis, the levels of ABA and GA1+3decreased sharply in the ears of “97J1”, while in the ears of “H8679” there was only a slight decrease in ABA, and even an increase in GA1+3. The ratio of isopentenyladenosine (iPA)/ABA and IAA/ABA in the ears of “97J1” increased sharply from antherlobe formation to meiosis, but changed only slightly in the ears of “H8679”. At antherlobe formation, IAA and GA1+3 levels were higher in the ears of “97J1”, but lower in the ears of “H8679”than in the leaves. At meiosis, ABA, GA1+3 and IAA levels in the “97J1” ears were much lower than in the leaves, but similar in “H8679”. These results indicated that the sharp decreases of ABA and GA1+3 in ears from antherlobe formation to meiosis and the lowest maintenance at meiosis may be favorable for development of fertile florets and enhancement of grain set in wheat.     

Effects of Boron Deficiency on Anther Development and Floret Fertility in Wheat (Triticum aestivum L. 'Wilgoyne')

Boron (B) deficiency limits reproductive growth more than vegetativegrowth in cereals such as wheat. The purpose of the presentstudy was to identify the critical stages of anther developmentof wheat (‘Wilgoyne’) during which B deficiencycauses a significant and irreversible decrease of floret fertilityin order to formulate timely measures for correcting or preventingthis problem. Withdrawing B from the rooting medium for 3 dbetween premeiotic interphase through meiosis to late tetrad,limited anther elongation and resulted in the loss of pollenviability. The negative effects of B withdrawal on anther lengthsuggest that the role of B in reproductive cell walls is similarto that in vegetative ones. The results indicated that as moreflorets reached meiosis within the period of B withdrawal, thelower was floret fertility and the number of grains set in awhole ear. For the whole ear, the critical period during whichB deficiency causes maximal and irreversible damage to floretfertility was about 7 d, extending from the early emergenceof the flag leaf to 2–3 d after its full emergence. Theresults suggest that there are two phases of pollen developmentsensitive to boron deficiency: the period from premeiotic interphasethrough meiosis to late tetrad was the most sensitive stageof microsporogenesis in wheat while the period from mitosis-Ito II during which starch accumulation occurred in pollen grainswas less sensitive. Copyright 2000 Annals of Botany Company Anther, boron (B) deficiency, floret fertility, Grain Set Index (GSI), pollen, wheat (‘Wilgoyne’)     

不同穗型小麦小花发育过程中幼穗和叶片内源激素水平的动态变化

以“９７鉴１”、“扬麦１５８”和“河南８６７９”分别代表特大穗型、大穗型和穗数型基因型材料,测定和分析了小麦（Ｔｒｉｔｉｃｕｍ ａｅｓｔｉｖｕｍ Ｌ．）从小花分化至抽穗期穗部和叶片的内源植物激素水平。幼穗中ＡＢＡ高峰出现在雌雄蕊分化期至药隔形成期之间,出现时间则随穗型增大而推迟。在减数分裂期前,特大穗型幼穗中内源激素水平特别是,ＡＢＡ和ＧＡ１＋３水平明显下降,但ＩＡＡ／ＡＢＡ及ｉＰＡ／ＡＢＡ的比值     

Photosynthesis and Transpiration in Leaves and Ears of Wheat and Barley Varieties

Carbon exchange rate (CER) and transpiration were measured inflag leaves, whole ears, glumes (referring to the total areaof glumes and lemmas) and awns, in six hexaploid spring wheats(Triticum aestivum L.), three cultivated tetraploid spring wheats(T. turgidum L.), four wild tetraploid wheats (T. dicoccoides),eight six-rowed barleys (Hordeum vulgare L.) and five two-rowedbarleys (H. vulgare L.). Differences between varieties and between species in total earCER and transpiration were associated largely with differencesin ear surface area rather than with rates per unit area. Ratesof CER and transpiration per unit area of ears were 40–80%of those of flag leaves, depending on the species. However, since ear surface area was greater than flag leaf areaby a factor of 1.1, 3.9, 5.5 and 4.4, in hexaploid wheat, tetraploidwheat, six-rowed barley, and two-rowed barley, respectively,total ear CER reached up to 90% of that of the flag leaf. The contribution of awns to total ear CER depended largely ontotal awn surface area per ear, rather than on CER per unitawn area. Awns contributed about 40–80% of total spikeCER, depending on the species, but only 10–20% of spiketranspiration. The disproportionately small contribution ofawns to ear transpiration was caused by the very low rate oftranspiration per unit area of awns. Thus, while transpirationratio (CER/transpiration) was about the same in flag leavesand glumes, it was higher by several orders of magnitude inthe awns. A large amount of awns in the ear is therefore a drought adaptiveattribute in these cereals, for which tetraploid wheat exceededhexaploid wheat and six-rowed barley exceeded two-rowed barley. Key words: Carbon exchange rate, Transpiration, Barley, Wheat     

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     

Photosynthesis of Ears and Flag Leaves of Wheat and Barley

Immediately after anthesis ears of spring wheat absorbed lessthan 0.5 mg CO₂, per hour in daylight and later evolved CO₂,in the light and in the dark. The rate of apparent photosynthesisof the combined flag-leaf lamina and sheath and peduncle (collectivelycalled flag leaf) of two spring wheat varieties, Atle and JufyI, was 3–4 mg per hour; the rates of the flag leaf andthe ear of two spring barleys, Plumage Archer and Proctor, wereeach about 1 mg per hour. The gas exchange of ears and flag leaves between ear emergenceand maturity accounted for most of the final grain dry weight.The CO₂, fixed by the wheat ear was equivalent to between 17and 30 per cent of the grain weight, but more than this waslost by respiration, so assimilation in the flag leaf was equivalentto 110–20 per cent of the final grain weight. In barley,photosynthesis in the flag leaf and the net CO₂ uptake by theear each provided about half of the carbohydrate in the grain. Barley ears photosynthesized more than wheat ears because oftheir greater surface, and flag leaves of wheat photosynthesizedmore than those of barley because they had more surface anda slightly greater rate of photosynthesis per dm².     

Influence of Vernalization and Photoperiod on Supernumerary Spikelet Expression in Wheat

Two tetraploid (Triticum turgidum L.emend gr. turgidum and gr.durum) and five hexaploid wheats (Triticum x aestivum L. emendgr. aestivum) with reported tendencies for ‘branched heads’(supernurnerary spikelets) exhibited variation in its expressionunder different vernalization photoperiod and temperature regimes. Two main types of supernumerary spikelets were identified, multiplesessile spikelets (MSS) with two or more complete spikeletsat a rachis node and indeterminate rachilla spikelets (IRS)with two to 13 spikelets on an extended rachilla. The degree of supernumerary spikelet expression in wheats withvernalization response differed from those without. Short photoperiods(9–14 h) both outdoors and in a glasshouse environment,were more conducive to supernumerary spikelet expression than24 h photoperiod in both environments. The 24 h photoperiodglasshouse environment (higher mean temperatures) was leastconducive to its expression except in lines with a strong vernalizationresponse. The high stability of supernumerary spikelet expression in certaingenotypes in the different environments indicated the feasibilityof incorporating this character in breeding and selecting commercialwheats to increase grain number per head. Triticum, wheat, ear-branching, supernumerary spikelets, vernalization, photoperiod, temperature     

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 of Flag Leaves and Ears of Wheat Hastened by Methyl Jasmonate

Treatment of flag leaves and ears of wheat plants with MJ (jasmonic acid methylester) (10⁻⁵ and 10⁻⁴ m) did not increase ethylene production, but it did accelerate senescence as indicated by the loss of chlorophyll. MJ also caused the closure of stomata, and consequently the rates of transpiration and photosynthesis decreased. Early maturity shortened the grain filling period, so the thousand grain weight was lower. Although ethylene elicited the same physiologic effects, the syndrome of senescence by MJ is independent of the former. We conclude that senescence and death in wheat are far from being elucidated; however, MJ and ethylene seem to participate in the phenomenon. Received July 10, 1997; accepted January 5, 1998     

The Chemical Induction of Supernumerary Shoots in the Developing Embryos of Wheat

The induction of seedlings with multiple shoots by the in vivo treatment of embryos of wheat by herbicides such as 2,4-D, 2,4,5-T, MCPA and TBA as well as mixtures of these substances was studied. Treatment during the first week of embryo development was necessary for the production of additional shoots in the seedling; later treatment gave rise to callused seedlings. Other seedlings such as oats, barley and rye, and other species of wheat were treated, but seedlings with extra shoots were not produced, though callused seedlings were recovered.     

Specificity of Pseudomonas Isolates on Healthy and Fusarium Head Blight-Infected Spikelets of Wheat Heads

The specificity of culturable bacteria on healthy and Fusarium head blight (FHB)-infected spikelets of wheat heads was investigated to find a candidate of biocontrol agents against FHB. The bacterial genus Pseudomonas was commonly isolated from the tissues, and phylogenetic analysis using 16S ribosomal RNA gene sequences of isolates of the genera revealed that particular phylogenetic groups in the genus specifically inhabited either healthy or infected spikelet tissues. The specificity of each group was suggested to be due to differences in the ability to form biofilms and colonize spikelet tissues; isolates originated from healthy spikelets formed biofilms on polyvinyl chloride microplate wells and highly colonized the spikelet tissues. Other bacterial groups obtained from FHB-infected spikelets less formed biofilms and attached with low densities on the spikelet tissues. Their colonization on the tissues, however, was promoted when co-inoculated with the causal pathogenic fungus, Fusarium graminearum, and several isolates were observed to smash the mycelia in vivo. Moreover, based on results of in vitro mycelial growth inhibition activity, the diseased tissue-originated isolates were verified to have a negative effect on the fungal growth. These results suggest that Pseudomonas isolates obtained from infected spikelet tissues were highly associated with the FHB pathogen and have potential as candidates for biological control against FHB.     

The Uptake of Surcrose and its Conversion to Starch in Detached Ears of Wheat

Detached ears of wheat were cultured on solutions of ¹⁴C-sucroseand the distribution of carbon-14 in the ear was followed. Within 8 h radioactive sucrose was found in all the tissuesof the cultured ear, and considerable amounts of carbon-14 hadaccumulated in other ethanol-soluble compounds. Carbon-14 accumulatedrapidly in the starch deposited in the endosperm, but littlewas found in the starch of the pericarp, or in other materialinsoluble in ethanol in the vegetative organs. During 48 h the specific radioactivity of the sucrose in theendosperm increased in a hyperbolic pattern and was equal tothat of the starch produced. Carbon-14 in the glucose and fructoseaccumulated more slowly and in a Linear fashion. Experiments with sucrose containing carbon-14 in both moietiesequally, or in the fructosyl moiety exclusively, confirmed thatboth moieties are converted into starch and at about the samerate. As sucrose in endosperm provided with asymmetric sucroseretained a considerable degree of asymmetry, it seems as thoughinversion is not a necessary step in the transport of sucroseinto the grain. In ears provided with ¹⁴C-sucrose at 30 mg ml^–1 the rateof accumulation of ¹⁴C-sucrose in the culm, rachis, and floralorgans was about 0.6 times the value at 50 mg ml^–1. However,in the sucrose of the endosperm, and in the starch depositedthere, the rates of accumulation of carbon-14 from both levelswere identical. This finding supports the concept that the transportof sucrose is limited during the final stages of its passageinto the grain.     

Localization and Distribution of the ATPase Activity and of the Passage of Material Transport in Spike-Stalk and Their Relations to the Development of Spikelets in Wheat

1. The ATPase activity in the spike-stalk cells of wheat was obviously localized at plasmallemma and the surface of cell wall bordering the intercellular spaces and their inclusions. The reactions of ATPase activity at chromatin and nucleoli were usually insignificant, and they were not found in vacuoles and other organelles (Figs. 1, 2, 3, 5, 6 and 7). 2. Three significant differences were observed between the middle part and the basal and upper part of the spike-stalk in wheat. (1) A large amount of inclusions were shown in the intercellular spaces of the middle part, and the high ATPase activity was seen at these inclusions (Figs. 6 and 7), but both they were seldom to be found in the intercellular spaces of the basal and upper part (Figs. 2 and 3). (2) The plasmodesmata of the middle part ceils was more than that of the basal and upper part ones (Figs. 1, 3 and 5). (3) In the middle part cells of spike-stalk, the cytoplasmic material was vigorously and actively transferred through the wall pores, and at the same time, the high ATPase activity was exhibited on the transferred cytoplasm (Figs. 4, 8 and 9). In addition, it was also observed that the cytoplasmic material entered into intercellular spaces from adjacent cells (Fig. 6). But it was hardly to see this phenomenon in the basal and upper part of spike-stalk. 3. It was discussed that the ATPase activity and the passage for material transport may play the role in transferring materials into the spike and they might be related to the development of the wheat spikelets.     

Changes in Respiration and Cyanide Sensitivity of the Barley Floret during Development and Maturation

The 6-week period of development and maturation of the barley (Hordeum vulgare L.) floret from anthesis to harvest is characterized by two phases: an early phase of rapid increase in respiration rate and dry weight, and a late phase during which respiration decreased rapidly whereas dry weight remained unchanged. Consumption of O₂ by the embryo changed little during the entire developmental period, whereas O₂ uptake by the endosperm and the lemma and palea decreased significantly during the late phase.