Studies in the Physiology of Flowering of Timothy (Phleum pratense L.): III. Effects of Shoot Age and Nitrogen Level on the Timing of Inflorescence Production

The influence of shoot age on flowering in S.48 timothy wasinvestigated in partially controlled conditions and outdoors.In artificial long days sensitivity to photo-induction increasedwith age of the shoot from soon after germination until aboutfive leaves were produced; no further increases in responseto photoperiod were evident in older plants. The minimum leafnumber to ear emergence in plants exposed to continuous lightfrom germination was nine; of these, five had been producedbefore spikelet initiation, while the remainder developed duringspike differentiation. In natural conditions, only main shoots on plants sown in Marchor earlier were sufficiently advanced to respond to the increasingnatural daylength and to produce ears in late June. In latersowings ear emergence became increasingly delayed until, inplants sown in June, some main shoots failed to develop inflorescences.Nitrogen deficiency further delayed or inhibited flowering inthese later-formed shoots. The minimum period of 10 weeks forinflorescence production in main shoots was attained in plantssown some weeks before midsummer. The equivalent period in subsidiarytillers arising in midsummer on the same plants decreased toless than 5 weeks. The implications of the results are discussedin relation to previous investigations in this species.     

Effect of plant population on ear differentiation and growth in maize

Maize was grown at 2.47, 3.70 and 4.94 plants m^-2. Ears were apparently initiated about 2 days later in the densest than in the least dense population, but rate of formation and final number of female floret primordia per row of the ear were similar in the two populations. In the densest population fewer primordia produced fully developed florets than in the least dense population and of those that were produced fewer extruded silks before pollen shedding ceased. Relative growth rates of dry mass and of length of the ear before flowering were unaffected by density, but with the apparent later start of ear growth in the densest population ears were lighter and shorter than in the least dense population. Number of florets per row of the ear at the time of flowering decreased slightly with increase in population. Number of kernels per row decreased with time after flowering, especially in the densest population. At final harvest size and mass per kernel, number of kernels per ear, length and circumference of the rachis, and space per kernel on the rachis all decreased with increase in population. Thinning the intermediate population at various times from shortly before flowering until near final harvest increased mass of grain and shoot, and number of kernels of the remaining plants at final harvest. The extent of the increases decreased progressively the later the thinning. Mass and volume per kernel were increased by thinnings done up to the time of flowering, but were unaffected by thinning from c. 16 days after flowering.     

Shoot growth dynamics and size-dependent shoot fate of a clonal plant,Festuca rubra, in a mountain grassland

The relation of the within-season and between-season patterns of shoot growth were compared in a clonal grass with long-lived shoots,Festuca rubra, in a mown mountain grassland. The growth rate of shoot length from spring to summer in a year was almost constant for each shoot irrespective of spring shoot length each year. The annual shoot growth rate from spring to spring was negatively correlated with the shoot length in the first spring. Shoots of different length and age therefore tended to converge over time to a population of identical shoot size, suggesting an equalizing effect of growth pattern on size structure. Shoot size (shoot length and number of leaves) influenced the fates of shoots. Larger shoots showed an increased incidence of both flowering and formation of intravaginal daughter shoots and a decreased incidence of death in the subsequent time period. The fates of shoots were independent of their age. Although the negatively size-dependent springto-spring annual shoot growth rate acted to decrease shoot size variation, the remaining variation within the shoot population was still sufficient to generate different fates of shoots. These fates were not related to the previous life history of individual shoots. There was a significantly positive effect of the shoot size at initiation on its life expectancy. This was mainly attributable to the positively size-dependent survival rate of shoots in the early stage (<1 year old) of shoot life history. Later on (> 1 year old), shoot size had little effect on the survival rate of shoots. Once small young shoots have survived this early stage (< 1 year old) in life history, they can grow vigorously, little affected by competition regardless of shoot size, and converge to a stable size structure of shoots of similar size. Only shoot size in the early stage ( < 1 year old) of life history is important for the persistence of a shoot population.     

Some treatments affecting growth substances in developing wheat ears

Cytokinin, auxin and gibberellin-like substances were bio-assayed in extracts from developing ears of wheat plants grown in various conditions. Changes in cytokinin activity along the ears may be related to the earlier flowering in the middle of the ear. Ears on the main stems of plants from which all the tillers had been removed contained less cytokinin than the main-stem ears of normal tillered plants. When grain development was stopped by preventing fertilization of the ovules the ear contained more cytokinin than normal ears. With de-tillered plants, removing flag leaves before anthesis increased cytokinin, gibberellin and auxin in the ears; later removal of flag leaves did not affect cytokinin but decreased gibberellin in the ears. Conversely, removing ears before anthesis did not affect cytokinin or auxin in the flag leaves, but their gibberellin was less than that of flag leaves on intact plants. Treatment of wheat ears with zeatin did not affect grain weight or number per ear which supports the conclusion that the growth substances in the ear may be adequate for normal grain growth.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE)

Individual tillers of timothy ( Phleum pratense ) growing from seed in pots were labelled and their life history studied over a period of two years.
The leaf axil position of the first-produced tiller depended on its time of origin. A minimum of five leaves on the parent axis was necessary before its appearance, and no tiller emerged from the axil of any of the three terminal leaves of a shoot.
Ear-bearing capacity was high in early-formed tillers, but decreased progressively with later dates of origin despite favourable day length; all tillers arising after the end of July failed to produce inflorescences. A greater proportion of tillers on the main stem formed ears than secondary tillers of the same age. Time of ear emergence varied only within a few days for all tillers arising before the end of April. The number of leaves formed up to ear emergence varied linearly with date of origin and tiller position, the lowest recorded number being six.
Dry weight and length of inflorescence as well as seed yield per tiller varied linearly with the date of origin of tillers. Primary tillers generally had longer and heavier inflorescences and produced more seed per stem than secondary tillers of the same age.
Tillers arising at all times were found to have a limited span of life, not normally exceeding just over a year. The mean age of ear-bearing tillers varied linearly with their date of origin. Death-rates of all tillers generally increased with their age. Relative death-rates of fertile and vegetative tillers rose rapidly in the autumn but decreased during winter and early spring. Vegetative tillers which elongated in the year of formation survived for a shorter time than equivalent tillers in which stem elongation was delayed until the following year.
The perennial habit of grasses is discussed in relation to tiller survival. Successive new formation and death of tillers keep the plant in a condition of continuous dynamic change.     

Relationship between reproductive behavior and new shoot development in 5-year-old branches of olive trees (Olea europaea L.)

The development of new shoots plays a central role in the complex interactions determining vegetative and reproductive growth in woody plants. To explore this role we evaluated the new shoots in the olive tree, Olea europaea L., and the effect of fruiting on new shoot growth and subsequent flowering. Five-year-old branches served as canopy subunits in order to obtain a global, whole-tree view of new shoot number, size and morphological origin. The non-bearing trees had many more shoots than the fruit-bearing trees, and a greater number of longer shoots. In both bearing conditions, however, the majority of shoots were less than 4 cm long, with shoots of progressively longer lengths present in successively decreasing frequencies. Six major shoot types were defined on the basis of apical or lateral bud origin and of parent shoot age. On fruit-bearing trees, the new shoots originated predominantly from the shoot apex, while on non-fruiting trees, they formed mainly from axillary buds, but in both cases, they tended to develop on younger parent shoots. The previous bearing condition of the tree was the main determinant for subsequent inflorescence development, which was independent of both shoot type and length. Thus, reproductive behavior strongly affected both the amount and type of new branching, but subsequent flowering level was more influenced by previous bearing than by the potential flowering sites on new shoots.     

Survival of Tillers and Distribution of Dry Matter between Ear and Shoot of Barley Varieties

Plants of Plumage Archer barley grown in pots produced moreshoots than did Proctor, but had fewer ears at maturity, becausesome shoots died about 7 days before ear emergence. The numberand position on the plant of the shoots that died were consistentfor particular growing conditions. Shoots that died were notalways the last to be produced. There were no consistent differencesin dry weight, leaf area, nitrogen content, or apex developmentbetween shoots that did and did not survive; nor did removingthree developing ears on older shoots affect the survival ofother shoots. Growth of Plumage Archer plants was not checkedby death of shoots; dry matter, nitrogen, and leaf area wereredistributed so that increase in surviving shoots compensatedfor losses in the dead shoots. The dry-weight ratio of ear to shoot was smaller for Proctorthan for Plumage Archer at ear emergence and anthesis, but atmaturity it was greater for Proctor, both when ears were shadedto prevent them photosynthesizing and when they were unshaded.The varietal difference in ear: shoot dry-weight ratio was reversedbecause the relative growth-rate of ears of Proctor was greaterand because more dry matter was lost from its shoots. Both changesmay have been caused by translocation to the ear of a greaterproportion of the assimilate from the shoots of Proctor thanof Plumage Archer. Leaves of Proctor appeared to be more efficientin producing dry matter for the ear than those of Plumage Archer.     

Photosynthesis and transpiration of the flag leaf in four spring-wheat cultivars

Co₂ exchange and transpiration rates of the flag leaves of four spring wheat (Triticum aestivum L.) cultivars, namely Glenlea, Neepawa, Opal and Kolibri, were compared using infra-red gas-analysis technique. The plants were grown in a controlled environment under an 18-h photoperiod, with day and night temperatures of 20 and 15° C, respectively. The time course of the CO₂-exchange rate (CER) of the flag leaf differed among cultivars. CER began to decrease rapidly some 2 weeks after ear emergence in Glenlea, Neepawa and Kolibri, but only after 4 weeks in Opal. The decline in CER of Glenlea, Neepawa and Opal was continuous throughout the period of grain development whereas in Kolibri CER was maintained at a constant level between the 4th and 6th weeks after ear emergence. The transpiration rates of the flag leaves of the 4 cultivars did not change markedly until 6–7 weeks after ear emergence, indicating that the reduction in CER was not primarily a response to increased stomatal resistance to the diffusion of CO₂. Removing the ear of the main shoot of intact plants failed to depress CER of the subtending flag leaf until 5 weeks after ear removal. Removing the ears of all the tillers of plants in which all but 3 tillers had been removed at ear emergence did not depress CER until 4 weeks after ear emergence, but removal of the ear of the main shoot of plants where all the tillers had been removed at ear emergence reduced the CER of the flag leaf 2 weeks after ear removal. Removal of tillers at ear emergence had a marked effect on the time course of CER and transpiration rates of the flag leaf. Both CER and transpiration rates of a 4-tiller plant were maintained at a higher level throughout ear development as compared to those of a one-tiller plant. The transpiration rate of the flag leaf of Glenlea increased during the later part of the life of the leaf even for one-tiller plants with no ear, indicating that such a stomatal response may be part of the normal course of leaf aging and not a response to a feedback stimulus from the ear.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE)

All tillers of timothy ( Phleum pratense ) growing in sand culture were labelled at weekly intervals. At each of eighteen dates between June and December a sample of plants was harvested, and dry weight and leaf area were determined for each group of tillers.
Tiller production was continuous throughout the experiment. Only those tillers which appeared before the end of July were able to form inflorescences, but among them the proportion of tillers with ears decreased with increasing lateness of origin.
Dry weight per plant increased rapidly for the first 14 weeks, and then did not change significantly until the end of the experiment. The flowering tillers lost dry weight after seed maturation; vegetative tillers, although smaller in size, continued to increase in number and weight. Dry weight per inflorescences was greatest in the main stem and decreased in subsequent tillers.
Leaf area per plant rose to a maximum soon after ear emergence and then declined, largely because the loss in leaf surface sustained by the flowering tillers was not offset by a corresponding increase in the vegetative tillers. Leaf area ratio decreased at first rapidly but later more gradually, while net assimilation rate was relatively high until maximum leaf area was almost attained, but then fell continuously to negligible values by October. Relative growth rate decreased throughout the experiment.
Initially the growth of the plant was largely determined by the flowering tillers in which, in close resemblance to the cereal plant, leaf area ratio and relative growth rate declined continuously. Tillers which did not flower were successively smaller in size and, because of their later origin, were exposed to less favourable environmental conditions by the time they had increased sufficiently in number to affect the growth of the plant.     

The Effect of Removing Florets or Shading the Ear of Barley on Production and Distribution of Dry Matter

Halving the number of florets per ear of barley shortly afterthe ears emerged increased the dry weight of the rest of theplant. The effect was greatest 17 days after the ears emerged,and until then the increase in weight of the shoot of the plantnearly equalled the decrease in ear weight. Later, when theshoots lost weight, the difference between shoots with 24 or12 florets per ear disappeared, presumably because all the extralabile carbohydrate was lost by respiration. Unshaded plantsor plants with shaded ears or shoots were affected similarly.Removing florets did not affect net assimilation rate althoughit slightly increased the rate of photosynthesis of the flag-leaflamina between 10 and 17 days after the ear emerged. The weight of the rest of the plant was decreased by shadingthe ear. Shading decreased ear weight at all samplings. Theeffect was small, especially when florets were removed. Thus the size of the sink provided by the ear for carbohydrateaffected the movement of carbohydrate from the shoot. This invalidatessome of the methods used to estimate photosynthesis by the ear.     

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.     

Assimilation and Translocation in Perennial Grasses

Single leaves, ears, or shoots of timothy (Phleum pratense L.)were exposed in light to ¹⁴CO₂, then left overnight, after whichthe plants were autoradiographed. The following conclusionswere drawn. Actively growing leaves retain all their assimilatesand import from older ones. Fully expanded leaves export butdo not import assimilates. Export begins before leaf expansionis complete, so import and export may for a time be simultaneous.Exports go at first to younger leaves and to roots, accumulatingat meristems. At later stages, exports move downwards ratherthan upwards. Buds and small tillers import from older shoots,but large tillers do not import from other shoots or exportto other large ones. Ears assimilate while still green, andimport assimilates from their associated flag leaves. Exportsfrom other leaves on flowering stems move downwards. These findings agree in general with those from other plants:they are discussed in relation to the vascular system of thegrass plant, and the need for further studies, particularlyquantitative ones, is emphasized.     

Uptake and distribution of nitrogen in wheat plants supplied with different amounts of nitrogen after stem elongation

The uptake of nitrogen and its distribution between shoots and between organs within shoots in wheat (Triticum aestivum) was studied from the start of stem elongation to 28 days after anthesis in a glasshouse experiment with eight nitrogen levels between 0·1 and 12·8 mequiv./litre. There was no net uptake of nitrogen in plants supplied with 0·8 mequiv./litre or less; with more nitrogen the absorption increased linearly. Twenty to 44% of the total plant nitrogen was absorbed after anthesis, this fraction increasing with nutrient supply. The nitrogen allotted to the main shoot decreased until the onset of anthesis and increased thereafter at the expense of the tillers, except with 12·8 mequiv./litre, where nitrogen percentage in the main shoot decreased also after anthesis. Raising nitrogen supply increased the proportion of plant nitrogen recovered in the tillers. Nitrogen accumulated in the ear after emergence and by the 28th day after anthesis it contained between 52% and 73% of the total plant nitrogen. The ear of the main shoot had a higher proportion of shoot nitrogen than that of the tillers. The fraction of ear nitrogen supplied by retranslocation decreased from almost 100% with 0·8 mequiv. N/litre or less to nil with 12·8 mequiv./litre. Increasing nitrogen application decreased the fraction of total nitrogen allocated to the ear.     

Tillers Do Not Influence Phytotoxicity of Imazamethabenz in Wild Oat (Avena fatua)

The response of wild oat to imazamethabenz varies with the growth stage, but the role of tillers in this regard is unclear. Removal of tillers at the three-leaf stage before spraying with imazamethabenz did not significantly affect the total shoot fresh weight measured 3 weeks later. The leaf area and dry weight of intact plants at the three-leaf stage were 17–21% greater than for plants with coleoptilar and first leaf main shoot tillers (T0 and T1) removed. The greater leaf area may have increased herbicide interception per plant. Similar fresh weight reductions in main shoot, total tillers, and total shoots were found whether imazamethabenz was applied to the plant at the two-leaf without tillers or the three-leaf with two tillers stage. Imazamethabenz applied only to the main shoot reduced total shoot dry weight more than an equivalent amount of imazamethabenz applied only to tiller T1 or applied over the whole shoot. Imazamethabenz had the least inhibitory effect on whole plant growth when applied only to T1. When ¹⁴C-herbicide was applied to the first main shoot leaf of plants at the three-leaf stage with two tillers, the ¹⁴C translocated 38% to roots, 33% to the main shoot, and nearly 30% to all tillers. When ¹⁴C-herbicide was applied to the first leaf of T1 then the ¹⁴C translocated 50% to T1, 25% to the main shoot, 20% to roots, and 5% to all other tillers. The translocation pattern and fresh weight values suggested that the presence of early tillers during herbicide application neither increased nor decreased imazamethabenz efficacy in wild oat. Received June 4, 1997; accepted June 5, 1997     

The effect of tiller removal and tiller defoliation on competition between the main shoot and tillers of spring barley

In glasshouse and field experiments the source-sink relations of the main shoot of plants of spring barley were modified by tiller removal and tiller defoliation. Decreasing competition by tiller removal promoted the growth of the residual main shoot and its component parts, and the earlier tillers were removed the greater was the effect. Stem dry weight was increased four-fold in the glasshouse by early tiller removal and was doubled in the field experiment. The grain yield of the main shoot ear was increased by 26 – 30% by tiller removal compared with tillering control plants and this was due to larger grains in all spikelet positions. On the other hand increasing competition by regular tiller defoliation had relatively little effect on the growth and development of the main shoot in the glasshouse study, but in the field the main shoot grain yield was reduced by 10% compared with the control. The main effect of tiller defoliation was on the development of tillers. In the glasshouse tillers survived repeated defoliation, continued to be produced, and the majority produced grain but with fewer and smaller grains per ear than in control plants. Tiller growth was supported by the import of assimilate from the main shoot and this was accompanied by an increase in the photosynthetic rate of the main shoot leaves. In the field all defoliated tillers died within 4 wk. These responses are discussed in terms of the physiological interrelations between the main shoot and tillers.     

Analysis of quantitative traits connected with seed weight and flowering terms in birdsfoot trefoil (Lotus corniculatus L.) plants

The nature of inheritance of characters connected with the seed size (seed mass) and quality of the shoots (germination energy, seed viability, general shoot length, shoot mass, dry weight of shoots) in Lotus corniculatus L. was analyzed. The investigations were carried out on plants of three varieties: the wild form from the Krasnodar Region and local forms MF1 and MF3. The correlation analysis was carried out. The pattern of inheritance in the terms of plant flowering and length of shoots were studied.     

A Developmental Pattern of Flowering in Colored <Emphasis Type="Italic">Zantedeschia</Emphasis> spp.: Effects of Bud Position and Gibberellin

The restricted flowering of colored cultivars ofZantedeschia is a consequence of developmental constraints imposed by apical dominance of the primary bud on secondary buds in the tuber, and by the sympodial growth of individual shoots. GA₃ enhances flowering inZantedeschia by increasing the number of flowering shoots per tuber and inflorescences per shoot. The effects of gibberellin on the pattern of flowering and on the developmental fate of differentiated inflorescences along the tuber axis and individual shoot axes were studied in GA₃ and Uniconazole-treated tubers. Inflorescence primordia and fully developed (emerged) floral stems produced during tuber storage and the plant growth period were recorded. Days to flowering, percent of flowering shoots and floral stem length decreased basipetally along the shoot and tuber axes. GA₃ prolonged the flowering period and increased both the number of flowering shoots per tuber and the differentiated inflorescences per shoot. Activated buds were GA₃ responsive regardless of meristem size or age. Uniconazole did not inhibit inflorescence differentiation but inhibited floral stem elongation. The results suggest that GA₃ has a dual action in the flowering process: induction of inflorescence differentiation and promotion of floral stem elongation. The flowering pattern could be a result of a gradient in the distribution of endogenous factors involved in inflorescence differentialtion (possibly GAs) and in floral stem growth. This gradient along the tuber and shoot axes is probably controlled by apical dominance of the primary bud. Online publication: 7 April 2005     

Effect of the timing of water deficit on growth, phenology and yield of pearl millet (Pennisetum glaucum (L.) R. Br.) grown in Sahelian conditions

Several studies conducted under high input conditions have indicatedlittle susceptibility of pearl millet to water deficit untillearly grain filling, because the losses in main shoot productionwere fully compensated by increased tiller fertility. The presentstudy assessed the impact of water deficits at three developmentstages: prior to flowering (S30), at the beginning of flowering(S45), and at the end of flowering (S60) in pearl millet grownin experimental conditions similar to Sahelian farming conditions.It included a control irrigation treatment simulating the naturaldistribution of rainfall throughout the cropping season. Bothbiomass production and grain yield were severely reduced byS30 and S45, while S60 had no effect. In S30 and S45, the floweringof tillers was delayed or totally inhibited. In both of thesetreatments, the low number of productive tillers did not compensatefor damage to panicle initiation and flowering of the main shoot.All treatments maintained green leaves on the main shoot duringthe grain filling period, and in S30 leaf growth recovered frommid-season drought. These results illustrate how pearl milletmostly escapes drought by matching its phenology to the meanrainfall distribution in the Sahel. In the case of mid-seasondrought, some late productive tillers and the maintenance ofgreen leaf biomass of the main shoots limited, but did not overcome,the yield losses. This study stresses the importance of agro-ecologicalconditions in control treatments, particularly the water regimeand crop density, when assessing crop drought resistance. Key words: Drought resistance strategy, experimental conditions, main shoots, tillers, Pennisetum glaucum     

Consequences of the Elimination of Old Leaves upon Spring Phenological Events and the New Leaves Nutrient Concentration in a Wintergreen Woody Species in the Southern Hemisphere

Previous studies analyzed the importance of old leaves conservancy for wintergreen species plant growth only after early spring old leaves elimination. However, carbon and nutrient resources for growth could have already been translocated from old leaves to shoots during autumn. In this work, the effect of old leaves absence on the leaf mass per area (LMA, g m⁻²) and nutrient concentration of new spring leaves, shoot growth, and flowering was studied in Aristotelia chilensis, an Andean Patagonic woody wintergreen species of Argentina. Plants were studied after autumn defoliation (AD) or late winter defoliation (WD) and results were compared to those of undamaged control plants (CO). The new leaves LMA and mineral nutrient (N, P, K, and Mg) concentration values did not decrease in AD or WD compared to CO plants. Conversely, CO plants showed higher flowering intensity and shoot lengthening compared to AD or WD plants. There were not remarkable differences regarding the defoliation time, though non-flowering shoots grew in a lesser degree than the flowering shoots in WD plants. It was concluded that A. chilensis old leaves cohort is an important source to shoot growth and flowering but their absence does not affect the new leaves structure or nutritional status from early spring in either AD or in WD plants. New leaves formation probably is guaranteed by resources (carbon and nutrients) previously stored in stems or even in the buds containing the preformed leaves since March, by the end of summer. Provided the availability of complete resources for the new leaf flush independently of the old leaves A. chilensis would restore the carbon balance as soon as possible to resume the growth of heterotrophic tissues at normal rates. Endogenous response to counterbalance the old leaves absence on non-flowering shoots was more effective when there was greater lag time between defoliation and shoot growth resume. Flowering and non-flowering shoots compete for the available resources when A. chilensis have not yet expanded leaves and shoots supporting reproductive structures were stronger sinks compared to non-flowering shoots in WD plants.     

Effectiveness of SC2053 as a chemical hybridizing agent for winter wheat

The use of chemical hybridizing agents (CHA) allows production of hybrid wheat seeds. We evaluated the effectiveness of a new CHA (SC2053) to induce male sterility on winter wheat in controlled growth conditions. CHA effectiveness was measured with the application of 4 doses (0–400–700–1000 g.ha^–1) at 7 stages. These stages were defined by the length of the main stem spike (1–4–7–11–15–20–40 mm). At heading, individual ears were isolated with a greaseproof paper bag. The seeds formed were counted on treated and control ears. The spikes' sterility was calculated three weeks after flowering. The sterility of the main stem's spike reached 95% to 100% for application of 700 g.ha^–1 and 1000 g.ha^–1 for main stem spike length of 7 mm to 20 mm. The effects of ear tillering (5 tillers per plant) on CHA effectiveness were also investigated. We observed a significant delay of ear development between the main stem and tillers so that complete sterilities were not reached for each dose. Since tillering in field conditions rarely exceeds 3 ears per plant, CHA effectiveness was studied on plants bearing 3 ears. The mean sterility of the first 3 ears was close to 100% for applications with 700 g.ha^–1 and 1000 g.ha^–1 at stages from 11 mm to 20 mm of main stem spike length.