Promotion of Flowering on Mature Picea sitchensis by Gibberellin and Environmental Treatments. The Influence of Timing and Hormonal Concentration

Male and female flowering were significantly promoted by gibberellin A_4/7+ A₃ mixtures applied beneath bark flaps on branches of 11-year-grafted mature scions of Picea sitchensis (Bong.) Carr. growing in the open in Scotland; 4 mg/ml gave more flowering than higher or lower concentrations of gibberellins and application in June was more effective than in July. Gibberellins also increased male flowering on mature scions when applied directly to the buds in a polythene house. A gibberellin A_4/7+ A₃ mixture produced greater numbers of strobili when applied to 2-year-grafted plants in May and June than when applied in July and August. Flowering of 5-year-grafted plants increased with increasing concentrations of a gibberellin A_4/7+ A₉ mixture up to 19.7 mg/ml above which flowering decreased. Modifying the environment by keeping in a polythene house enhanced the combined male and female flowering of 5-year-grafted mature scions above the amount on control plants outside, the effect being more marked in June and July than in August and September. Larger numbers of gibberellin applications or longer total lengths of time in the polythene house gave significantly greater amounts of combined male and female strobili.     

Promotion of Female Strobili Flowering and Seed Production in two Japanese Pine Species by 6-Benzylaminopurine (BAP) Paste Application in a Field Seed Orchard

Treatment with 6-benzylaminopurine (BAP) to promote the flowering of female strobili and to increase seed production in a seed orchard was examined at different times of conebud differentiation for Japanese red pine (Pinus densiflora) and Japanese black pine (P. thunbergii), from late August to mid-September 1998. BAP was mixed with lanolin and Vaseline to prepare a paste (BAP, 2000 mg/L), which was used for a terminal bud treatment and also applied in a girdling treatment. The BAP treatment in early September transformed lateral buds on lower branches that would have normally differentiated as male strobili into female strobili. The highest percentages of female flowering branches were 21.1% (red) and 40.5% (black) produced by the terminal bud treatment, while control branch percentages were 5.0% and 7.1%, respectively. Girdling combined with BAP treatment further promoted female flowering branches to 32.3% (red) and 64.3% (black), while those of controls were 8.3% and 9.9%, respectively. Moreover, the numbers of female strobili per female flowering branch were 6 to 11-fold greater than controls. Mature seed cones were harvested in October two years later from which full seeds with a high germination ratio were recovered. Use of BAP treatment is thus highly effective for enhancing seed production in seed orchards of these two pine species.     

Promotion of flowering in the Pinaceae by gibberellins

Significant female flowering of 6- to 11-year-old seedlings and grafted ramets of sexually mature scions of lodgepole pine (Pinus contorta Dougl.) was promoted by both topical and spray applications of a gibberellin (GA) A_4/7 mixture (1.6 to c. 5 mg per plant in total) during that period (June to September) when sexual differentiation of lateral primordia would be expected to take place. Girdling was used in most experiments to enhance the GA_4/7 effect, as was the auxin, naphthaleneacetic acid (NAA). Average frequency of flowering branches on treated plants over all experiments ranged from 27 to 59% (control ranged from 0 to 36%) and average number of female strobili was increased from 2- to 6-fold by growth regulator treatment, relative to controls. Within an experiment, clonal or family frequency of flowering for treated plants ranged from 11 to 67% (controls were 0 to 28%), and number of female strobili was increased from 2- to 14-fold by growth regulator treatment, relative to controls. Movement of the flowering stimulus from the point of application was apparent in several experiments, the response in adjacent branches being correlated positively with increasing dosage of GA_4/7. Significant male flowering occurred only in one experiment, girdling and GA_4/7 treatment being promotive factors. The use of spray applications of GA_4/7+ NAA is warranted to induce early and enhanced flowering in lodgepole pine seedlings and vegetative propagules for genetic improvement programs.     

Studies of Growth and Flowering in Picea sitchensis (Bong.) Carr. 1. Effects of Growth Regulator Applications to Mature Scions on Seedling Rootstocks

Effects of growth regulator applications on the flowering of5-years-grafted mature scions of Picea sitchensis (Bong.) Carr.were assessed. Growth regulators were applied to the bud surfacein droplets of ethanol during July, August and September ina polythene house or glasshouse. A mixture of gibberellins A₄and A₇ applied alone, and in combination with gibberellins A₃and A₅, significantly increased numbers by up to seven timesfor male and by up to eight times for female strobili, gibberellinA, gave relatively the strongest response, and gibberellin A₄was inactive. Phosphon D and abscisic acid each reversed thepromotion of flowering by gibberellins, whilst kinetin and N,N-diphenylureahad no effect. The number of female strobili was negativelycorrelated with vegetative shoot length in the year after treatment. Under field conditions hormones were applied in July and Augustunder flaps of bark on the branches of 10-years-grafted maturescions. Gibberellin applications caused a 5-fold increase inflowering and N⁶-benzyladenine further increased the response.Naphth-lyl-acetic acid reduced female and increased male flowering.Bark removal near the base of the branch further enhanced hormone-inducedstrobilus production. The usefulness of these findings for thebreeding of Picea sitchensis is discussed.     

Maturation in larch : I. Effect of age on shoot growth, foliar characteristics, and DNA methylation

The time course of maturation in eastern larch (Larix laricina [Du Roi] K. Koch) was examined by grafting scions from trees of different ages onto 2-year-old root stock and following scion development for several years. Height, diameter, foliar chlorophyll content, and rooting ability of scion-derived cuttings all varied linearly as a function of log₁₀ age. Chlorophyll content (milligrams per gram of dry weight) increased while height, diameter, and ability to root decreased with age (P < 0.01). The tendency toward orthotropic growth and branch formation per centimeter of main stem decreased abruptly between age 1 and 5 years (P < 0.01). Total chlorophyll content of both long and short shoot foliage increased by 30 to 50% with increasing age, but the chlorophyll a/b ratio did not change. Also, juvenile long shoot needles were significantly longer than mature (P < 0.01). Surprisingly, the juvenile scions produced more total strobili over two successive years, but the mature scions produced a significantly higher proportion of male strobili (P < 0.001 year 1; P < 0.02 year 2). The age-related changes in foliar traits were not associated with changes in DNA methylation between juvenile and mature scions. Using HPLC, we found that 20% of foliar DNA cytosine residues were methylated in both scion types.     

The Cold Awakening of <Emphasis Type="Italic">Doritaenopsis</Emphasis> ‘Tinny Tender’ Orchid Flowers: The Role of Leaves in Cold-induced Bud Dormancy Release

Massive flowering of tropical Phalaenopsis orchids is coordinated by the cold-induced release of reproductive bud dormancy. Light and temperature are the two key factors integrated by the dormancy mechanism to both stop and reactivate the meristem development of many other angiosperm species, including fruit trees and ornamental plants. It is well established that leaves and roots play a major role in inducing flower development; however, currently, knowledge of molecular events associated with reproductive bud dormancy release in organs other than the bud is limited. Using differential gene expression, we have shown that the leaves of a hybrid of Phalaenopsis species, Doritaenopsis ‘Tinny Tender’, undergo major metabolic modifications. These changes result in the production of sucrose and amino acids, both of which can sustain bud outgrowth, and auxin and ethylene, which may play important roles in awaking the dormant buds. Intake of abscisic acid and synthesis of the hormone jasmonate may also explain the inhibition of vegetative growth that coincides with bud growth. Interestingly, many genes that were upregulated by cold treatment are homologous for genes involved in flower induction and vernalization in Arabidopsis, indicating that processes regulating flowering induction and those regulating reproductive bud dormancy release may use similar pathways and effector molecules.     

温度对叶用莴苣春化的影响

为了探讨温度对叶用莴苣春化的影响,进而明确其春化类型,本试验选用两个易抽薹叶用莴苣品种GB-30和GB-31为试材,分别进行20 d的4、20和25 ℃处理,然后均进行高温胁迫促进抽薹,采用石蜡切片观察茎尖花芽分化进程,并结合后期抽薹开花情况来判断是否进行春化,并确定其春化类型.结果表明:两个品种在前期经过3种温度处理后,在高温胁迫第8天时均有不同程度的抽薹.两个品种不同温度处理下花芽均进行了分化,前期4 ℃处理并没有使其花芽分化明显提前,后期的高温胁迫可以明显促进花芽分化进程.两品种间在不同温度处理下完成各发育阶段所需天数存在差异,两品种各处理从催芽到开花以及从高温到开花的有效积温也存在差异.前期未经低温处理的叶用莴苣也进入花芽分化、抽薹、现蕾及开花阶段,表明叶用莴苣为非低温春化型植物,后期高温处理能明显促进其抽薹开花,而且从催芽开始只有有效积温达到约2500 ℃·d才能开花.     

Dormancy and Flowering Are Regulated by the Reciprocal Interaction Between Cytokinin and Gibberellin in <Emphasis Type="Italic">Zantedeschia</Emphasis>

Floral productivity of Zantedeschia is dependent on the conversion rate of buds to shoots, which is controlled by varying intensities of para- (apical dominance), endo- (dormancy), and ecodormancy. We present evidence of cross-talk between cytokinin and gibberellin in their complementary roles to alleviate bud dormancy and enhance flowering in a perennial geophyte. We assessed the impact of cytokinin and gibberellin, applied alone and in sequential combinations, on bud fate during three phases along the ontogeny of growth, which coincide with the progressive transition of buds from apical dominance to dormancy. Given that cytokinin can stimulate branching and gibberellin can induce flowering in Zantedeschia, we measured these phenotypic responses as parameters of bud commitment. The efficacy of cytokinin alone to stimulate branching declined with the transition to dormancy (phase 1 = 3.8 ± 0.2 shoots; phase 3 = 1.0 ± 0.3 shoots). To sustain branching during this transition, a sequential application of gibberellin was necessary. Gibberellin alone failed to stimulate branching. The efficacy of gibberellin alone to stimulate flowering diminished with the transition to dormancy. Any flowering during this transition occurred only after the sequential application of cytokinin. Cytokinin alone failed to stimulate flowering. Alleviating bud dormancy and enhancing flowering in Zantedeschia, achieved by the reciprocal cross-talk between cytokinin and gibberellin, contributes to the pool of evidence drawing common mechanisms between dormancy and flowering and may have commercial implications.     

Studies of Growth and Flowering in Picea sitchensis (Bong.) Carr. 2. Initiation and Development of Male, Female and Vegetative Buds

Vegetative shoots from the base of the crown, and from partsof the tree likely to form male or female buds, were collectedfrom 40–years–old trees of Picea sitchensis (Bong.)Carr. throughout the 1973–4 annual growth cycle. The morphologyand growth rates of the terminal buds on these shoots were assessed. Bud scale primordia were formed most quickly in the female position,at an intermediate rate in the male position and most slowlyin the basal vegetative position during April, May and June.In July and early August the apical meristems swelled to formdomes and continued to grow at the same relative rates in themale, female and basal vegetative positions. Reproductive budswere first morphologically distinct in late August and sporangiaappeared in October. Dormancy, defined by the pause in apicalvolume increase, extended from mid-October to mid–March.Young strobili grew much faster than basal vegetative shootsof the same age between mid–March and bud burst in lateApril. Throughout the growth cycle, external changes in budsize reflected changes in size of the apical meristem, youngstrobihis or young vegetative shoot inside the bud. It is proposed that the rate of growth of an apical meristemmay be causally related to the type of bud which subsequentlydevelops from it. Sitka spruce, Picea sitchensis, bud development, morphology, growth of apical dome, flowering     

Chemical Factors Controlling Floral Bud Formation of Torenia Stem Segments Cultured in Vitro II. Effects of Growth Regulators

Correlative effects between growth regulators added to a mediumand different physiological states of explants on adventitiousbud formation and flowering were investigated using Toreniastem segments cultured in vitro. Indoleacetic acid stimulatedfloral bud formation and its development in explants taken fromreproductive plants. These stimulative effects were clearlyseen in explants taken from plants in which flower abscissionwas taking place, but insignificant when explants were preparedfrom younger materials. Abscisic acid acted in a reverse wayto auxin, greatly promoting floral bud initiation and floweringof originally vegetative explnts. Zeatin at a concentrationof 1 mg/liter inhibited floral bud formation, and at a low concentrationsit was generally ineffective. However, floral bud formationand flowering of explants taken either from basal parts of stemsor from 18- to 20-week-old plants were promoted by zeatin treatment.The action of gibberellic acid seemed rather indirect: at aconcentration of 0.01 mg/liter, it generally stimulated floralbud formation but at a concentration of 1 mg/liter, it was ofteninhibitory. (Received January 17, 1981; Accepted February 21, 1981)     

Effects of Oxygen and Respiratory Inhibitors on Induction and Release of Dormancy in Aerial Tubers of Begonia evansiana

Effects of O₂ and some respiratory inhibitors on the induction and release of bud dormancy were examined with the aerial tubers of different ages of Begonia evansiana Andr. Oxygen was needed not only for tuber sprouting but also during the chilling process at 2 to 5 C to break tuber dormancy. If the mature tubers were exposed to blue light during the chilling period, their dormancy was strikingly released even by the chilling given under an O₂ concentration as low as 3%. Blue light pretreatment promoted photo-sprouting of immature tubers only when given under lower O₂ concentrations. On the other hand, red light became effective in inducing dormancy in the immature tubers and in prolonging dormancy in the mature tubers as O₂ tension was increased. This was also the case with the induction of dormancy in the immature tubers by exposing them to a lower temperature (17 C) in the dark. The development of dormancy was suppressed by 2,4-dinitrophenol, p-nitrophenol, and sodium azide.     

Shoot growth components and flowering phenology in grafted<Emphasis Type="Italic"> Pinus halepensis</Emphasis> Mill.

Shoot elongation and flowering were assessed for a season (January–November) in 25 grafts from five clones of Pinus halepensis growing in a seed orchard. A co-dominant shoot from the upper crown and a dominated, low shoot were measured from each ramet. Upper shoots elongated continuously from a variable onset date between January and March and followed a logistic function against Julian day and a Gompertz function against heat sum above 0°C. Three to seven (averaging five) successive cycles were formed through the growing season; usually, two of them were preformed in the terminal bud (spring cycles) and one to four were neo-formed, summer cycles. The number of summer cycles and their contribution to the annual shoot growth were the only variables with a significant clonal influence. Ovulate strobili appeared from February to April and in October. Some ramets showing two female flowering cycles in the same shoot were observed. Lower shoots, bearing pollinate strobili always displayed a single spring cycle preformed in the winter bud.     

Low temperature influence on flowering in Citrus. The separation of inductive and bud dormancy releasing effects

The flowering response of Owari Satsuma mandarin ( Citrus unshiu Marc) to low temperature treatments has been determined using potted trees and in vitro bud cultures. In potted trees the chilling treatments released bud dormancy and enhanced both sprouting and flowering, but these two responses could not be separated. However, bud cultures showed no dormancy, and a specific effect of low temperature on flower induction was demonstrated. Low temperature appears to have a dual effect, releasing bud dormancy and inducing flowering. Potential flower buds have a deeper dormancy than vegetative buds, and the first stages of flower initiation seem to occur before the winter rest period.     

不同激素对锦绣杜鹃的催花作用

在加温、补光条件下,研究赤霉素(GA3)、萘乙酸(NAA)和吲哚乙酸(IAA)处理对锦绣杜鹃花蕾发育、开花期及开花质量等的影响。结果表明:锦绣杜鹃花蕾经0～3000mg.L-1GA3涂抹后,其催花效果随处理浓度增大呈先升后降趋势,并于2000mg.L-1浓度下达到最佳,表现在花蕾发育快、开花期提前及开花质量高等;而0～3000mg.L-1NAA和IAA对锦绣杜鹃催花效果均不明显。锦绣杜鹃花蕾大小(包括长度和宽度)与开花期提前时间和花径之间均呈极显著正相关,与花蕾期则呈极显著负相关;花蕾期与开花期提前时间之间呈极显著负相关(P<0.01)。     

Mapping of Candidate Genes Involved in Bud Dormancy and Flowering Time in Sweet Cherry (Prunus avium)

The timing of flowering in perennial plants is crucial for their survival in temperate climates and is regulated by the duration of bud dormancy. Bud dormancy release and bud break depend on the perception of cumulative chilling during endodormancy and heat during the bud development. The objectives of this work were to identify candidate genes involved in dormancy and flowering processes in sweet cherry, their mapping in two mapping progenies ‘Regina’ × ‘Garnet’ and ‘Regina’ × ‘Lapins’, and to select those candidate genes which co-localized with quantitative trait loci (QTLs) associated with temperature requirements for bud dormancy release and flowering. Based on available data on flowering processes in various species, a list of 79 candidate genes was established. The peach and sweet cherry orthologs were identified and primers were designed to amplify sweet cherry candidate gene fragments. Based on the amplified sequences of the three parents of the mapping progenies, SNPs segregations in the progenies were identified. Thirty five candidate genes were genetically mapped in at least one of the two progenies and all were in silico mapped. Co-localization between candidate genes and QTLs associated with temperature requirements and flowering date were identified for the first time in sweet cherry. The allelic composition of the candidate genes located in the major QTL for heat requirements and flowering date located on linkage group 4 have a significant effect on these two traits indicating their potential use for breeding programs in sweet cherry to select new varieties adapted to putative future climatic conditions.     

The Morphogenesis of Apple Buds: III. The Inception of Flowers

The early stages in the change from vegetative to reproductivedevelopment of apple spur terminal buds were followed by dissectionof buds from untreated trees, and from trees defoliated at differenttimes in the season. A change in the development of the leafprimordia occurred when there were approximately eight in thebud. This was followed by the development of bracts, which appearedto be necessary for the formation of actual flower parts. Leafprimordia tend to inhibit this process. Whereas their effectupon the apical meristem was subsequently reduced by the formationof bracts, so that eventually a terminal flower formed, theireffect upon the lower lateral meristems was unaltered. Thesemeristems therefore remained in a vegetative state. In addition to the number of leaf primordia in the bud, thedegree of dormancy may be an important factor in determiningthe onset of flowering. Since the number of leaf primordia invegetative buds at the end of the season is eight, the spatialdistribution of primordia on the main axis of the bud and theirvascular connexions might have a decisive effect on bud development.This was related to the effect of older primordia in the budupon the development of younger ones. In buds in which theseolder primordia were inhibited by foliage, etc., i.e. thosewith a long plastochrone, no effects were observed upon thedevelopment of younger primordia and the buds remained vegetative. Whilst correlative inhibition of buds thus affected their abilityto form flowers, there is no evidence of a critical leaf areafor flowering. Flowering in apple buds is more likely to bedue to the removal of factors inhibiting reproductive developmentthan to the synthesis of a specific flower inducing substanceas such.     

Levels and Aggregation of Ribosomes during Dormancy and Dormancy Break of Peach Flower Buds

The level of ribosomes and their aggregation state (monosomes-polysomes) have been investigated in relation to dormancy in peach flower buds (Prunus persica). The ribosomes were analysed by a linear sucrose gradient. The ribosomal content hardly changes in the bud before dormancy and until dormancy breaks; on the other hand after this period there is a remarkable increase in the ribosomal level strictly related to the increase in bud weight. The ribosomal content reaches the highest value at flowering. During dormancy break the largest portion of ribosomes is localized in flower primordia, not in scales. Polysomes, which are always present in buds, remain constant during dormancy (35%), increase at dormancy breaking, reaching the highest level at flowering.