Cytokinin/Auxin Control of Apical Dominance in Ipomoea nil

Although the concept of apical dominance control by the ratioof cytokinin to auxin is not new, recent experimentation withtransgenic plants has given this concept renewed attention.In the present study, it has been demonstrated that cytokinintreatments can partially reverse the inhibitory effect of auxinon lateral bud outgrowth in intact shoots of Ipomoea nil. Althoughless conclusive, this also appeared to occur in buds of isolatednodes. Auxin inhibited lateral bud outgrowth when applied eitherto the top of the stump of the decapitated shoot or directlyto the bud itself. However, the fact that cytokinin promotiveeffects on bud outgrowth are known to occur when cytokinin isapplied directly to the bud suggests different transport tissuesand/or sites of action for the two hormones. Cytokinin antagonistswere shown in some experiments to have a synergistic effectwith benzyladenine on the promotion of bud outgrowth. If theratio of cytokinin to auxin does control apical dominance, thenthe next critical question is how do these hormones interactin this correlative process? The hypothesis that shoot-derivedauxin inhibits lateral bud outgrowth indirectly by depletingcytokinin content in the shoots via inhibition of its productionin the roots was not supported in the present study which demonstratedthat the repressibility of lateral bud outgrowth by auxin treatmentsat various positions on the shoot was not correlated with proximityto the roots but rather with proximity to the buds. Resultsalso suggested that auxin in subtending mature leaves as wellas that in the shoot apex and adjacent small leaves may contributeto the apical dominance of a shoot. (Received September 24, 1996; Accepted March 16, 1997)     

The gravity-regulated growth of axillary buds is mediated by a mechanism different from decapitation-induced release

When the upper part of the main shoot of the Japanese morning glory (Pharbitis nil or Ipomoea nil) is bent down, the axillary bud situated on the uppermost node of the bending region is released from apical dominance and elongates. Here, we demonstrate that this release of axillary buds from apical dominance is gravity regulated. We utilized two agravitropic mutants of morning glory defective in gravisensing cell differentiation, weeping (we) and weeping2 (we2). Bending the main shoots of either we or we2 plants resulted in minimal elongation of their axillary buds. This aberration was genetically linked to the agravitropism phenotype of the mutants, which implied that shoot bending-induced release from apical dominance required gravisensing cells. Previous studies have shown that basipetal translocation of auxin from the apical bud inhibits axillary bud growth, whereas cytokinin promotes axillary bud outgrowth. We therefore compared the roles of auxin and cytokinin in bending- or decapitation-induced axillary bud growth. In the wild-type and we plants, decapitation increased cytokinin levels and reduced auxin response. In contrast, shoot bending did not cause significant changes in either cytokinin level or auxin response, suggesting that the mechanisms underlying gravity- and decapitation-regulated release from apical dominance are distinct and unique.     

Apical Dominanace in Xanthium strumarium: A DISCUSSION IN RELATION TO CURRENT HYPOTHESES OF CORRELATIVE INHIBITION

The influence of the spectral distribution of illumination onthe gibberellin, cytokinin, auxin, and abscisic acid levelsand the correlation with the degree of branching in Xanthiumstrumarium is presented and discussed. Gibberellins do not appearto play a major role in apical dominance but may be importantfor bud extension following the initial release from dominance.The cytokinin level was much higher in inhibited buds than inreleased buds. It is suggested that the cytokinins present wereprobably not able to participate in bud growth because of anauxin-induced accumulation of abscisic acid in the buds themselves.The concentration of abscisic acid as measured by bioassay andgas-liquid chromatography was between 50 and 250 times thatoccurring in all other plants parts examined. This level felldramatically following release from apical dominance by decapitation.The results are discussed in relation to current hypothesesof apical dominance.     

Early Cytological and Biochemical Events Induced by a 6-Benzylaminopurine Application on Inhibited Axillary Buds of Cicer arietinum Plants

Axillary buds of Cicer arietinum L. plants are released fromapical dominance by a direct 6-benzylaminopurine application.The cytokinin treatment induces a sequence of events, some featuresof which are described. An exponential increase of bud dry weightwas measured as soon as the sixth hour from the treatment andstarch accumulation was observed during the first 4 h. An importantincrease in uridine incorporation into RNA occurred 90 min afterapplication of the cytokinin. Cell division was stimulated veryearly while the increase of bud-DNA content was delayed. Theimplications of these results are discussed in relation to themechanisms of cytokinin action in the release of axillary budsfrom apical dominance.     

Roles for Auxin, Cytokinin, and Strigolactone in Regulating Shoot Branching

Many processes have been described in the control of shoot branching. Apical dominance is defined as the control exerted by the shoot tip on the outgrowth of axillary buds, whereas correlative inhibition includes the suppression of growth by other growing buds or shoots. The level, signaling, and/or flow of the plant hormone auxin in stems and buds is thought to be involved in these processes. In addition, RAMOSUS (RMS) branching genes in pea (Pisum sativum) control the synthesis and perception of a long-distance inhibitory branching signal produced in the stem and roots, a strigolactone or product. Auxin treatment affects the expression of RMS genes, but it is unclear whether the RMS network can regulate branching independently of auxin. Here, we explore whether apical dominance and correlative inhibition show independent or additive effects in rms mutant plants. Bud outgrowth and branch lengths are enhanced in decapitated and stem-girdled rms mutants compared with intact control plants. This may relate to an RMS-independent induction of axillary bud outgrowth by these treatments. Correlative inhibition was also apparent in rms mutant plants, again indicating an RMS-independent component. Treatments giving reductions in RMS1 and RMS5 gene expression, auxin transport, and auxin level in the main stem were not always sufficient to promote bud outgrowth. We suggest that this may relate to a failure to induce the expression of cytokinin biosynthesis genes, which always correlated with bud outgrowth in our treatments. We present a new model that accounts for apical dominance, correlative inhibition, RMS gene action, and auxin and cytokinin and their interactions in controlling the progression of buds through different control points from dormancy to sustained growth.     

Hormones and Apical Dominance in the Fern Davallia

Lateral buds of the fern Davallia trichomanoides are releasedfrom inhibition by the removal of the main shoot apex. However,auxin is not capable of substituting for the apex in decapitatedshoots nor can auxin in shoot tips be detected by bioassay orextraction and chromatography. Expanding leaves of this speciescontain auxin, but these organs are not responsible for inhibitionof lateral bud growth. The response of lateral buds to an exogenouslyapplied cytokinin does not result in initial bud break. It isconcluded that the hormonal factors known to govern apical dominancein seed plants are not responsible for the regulation of differentialbud expansion in this fern.     

Endogenous Abscisic Acid in torosa-2 Mutant Tomato Plant

Analyses of abscisic acid, phaseic acid and dihydrophaseic acidby high performance liquid chromatography were carried out onto-2 tomato mutant plant, which has strong apical dominanceexpression. Significant differences in comparison to the normalplant were found only at 20 days after sowing. Of particularinterest for the to-2 phenotype trait was the high quantitiesof these substances in the roots of this mutant. (Received April 20, 1984; Accepted November 21, 1984)     

Proteomic Analysis Points to a Role for RAD23 in Apical Dominance in Pinus sylvestris var. Mongolica

In higher plants, apical dominance is the phenomenon whereby the main central stem is dominant over other side stems. In the present study, the natural loss of apical dominance in a Pinus sylvestris var. mongolica mutant was characterized. The mutant lacked an obvious trunk, but exhibited a large number of side stems, short internodes and leaves, and dwarfism. Cell biological analyses indicated that the meristem was changed dramatically in the mutant, with irregular cell arrangements, and no obvious layers of meristem and mature tissue. The loss of apical dominance in the mutant plant was accompanied by a significant decrease in auxin (A) and cytokinin (C) content compared to wild type, and increased C/A values. Proteomic analysis of the apical buds using 2DE-ESI-MS/MS at different stages of development (including dormancy, bud break, 10 days after bud break, and 15 days after bud break) revealed 49 proteins that were significantly changed in abundance compared to wild type. These proteins were involved in stress response, energy production and conversion, post-translation modification and amino acid transport and metabolism, as well as many other complex biological processes. Among the identified proteins, Rad23, an abscisic acid- and stress-induced protein, and cytoplasmic aconitate hydratase likely participate in ABA responses, Fe²⁺ ion homeostasis and the ubiquitin/26S proteasome pathway to regulate apical dominance of buds. Rad23, in particular, appears likely to play an important role in the complicated mechanism of apical dominance of Pinus sylvestris var. mongolica. This work provides important insights into this process as well as forming a basis for the further study of plant apical dominance.     

The Presentation Time for Shoot Inversion Release of Apical Dominance in Pharbitis nil

The presentation time for shoot inversion release of apicaldominance in Pharbitis nil is between 1 and 1.5 d. Five to 6d of shoot inversion are required for persistent outgrowth ofthe highest lateral bud. Pharbitis nil, apical dominance, shoot inversion, lateral bud growth, presentation time     

Axillary Bud Inhibition Induced by Young Leaves or Bract in Euphorbia pulcherrima Willd

In plants held under long days in the vegetative stage, youngexpanding leaves of poinsettia (Euphorbia pulcherrima Willd.‘Brilliant Diamond’) are the main source of axillarybud inhibition, while the apical bud, which includes the meristem,primordial leaves and small unfolded leaves, is a secondaryinhibition source. Removal of these expanding leaves resultedin rapid release and growth of axillary buds. Decapitation ofthe apical bud resulted in delayed axillary bud release. Inreproductive plants kept in short days, the pigmented bractsare the primary source of axillary bud inhibition and the cyathiaare the secondary source. Applications of NAA —substitutedfor both young leaves and bract inhibition — maintainedapical dominance. The concentration of endogenous auxin washighest in the apical bud. However, when calculated on wholeorgan basis the auxin level was greater in young developingvegetative leaves and in reproductive bracts than in the apicalbud. Euphorbia pulcherrima Willd, apical bud, apical dominance, auxin, correlative inhibition, cyathia, poinsettia, IAA, NAA     

Differential Responses of Buds along the Shoot to Factors Involved in Apical Dominance

Intact and decapitated 6-node shoots of Hygrophila sp. weregrown aseptically immersed in liquid half-strength Knop's solutionwith microelements and 2% (w/v) sucrose (control medium), andin medium with 0.1 mg l^–1 benzyladenine (BA). In intactshoots grown in control medium apical dominance suppressed outgrowthof the lateral buds; in decapitated shoots buds grew out atseveral of the most apical nodes, increasing in size acropetally.There was a lag in outgrowth of the bud at the most apical node,attributable to its initially smaller size. Lateral shoots grewout first at basal nodes of intact shoots in BA medium, decreasingin size acropetally; in decapitated shoots in BA medium lateralshoots of approximately equal size grew out at all nodes. Differentialeffects of decapitation and cytokinin treatment on lateral shootoutgrowth along the shoot could be interpreted by postulatinga basipetally decreasing gradient of endogenous auxin concentrationin the intact shoot. Application of 20 mg l^–1 indoleaceticacid (IAA) in agar to decapitated shoots completely preventedbud outgrowth for at least 7 d in control medium, inhibitingit thereafter, and inhibited bud outgrowth in BA medium, thussupporting the hypothesis. Comparison of lateral shoot outgrowthin whole decapitated shoots and severed decapitated shoots (isolatednodes) lent no support to the alternative hypothesis that theremight be an acropetally decreasing concentration gradient ofa bud-promoting substance in the intact shoot, and demonstratedmuch greater lateral shoot growth in isolated nodes. The resultsemphasize important correlative relationships between the partsof a shoot with several nodes.     

Topophysis affects the Potential of Axillary Bud Growth, Fresh Biomass Accumulation and Specific Fresh Weight in Single-stem Roses (Rosa hybrida L.)

Topophysis, the effect on growth and differentiation of positionof axillary buds along the shoot, was studied by propagatingfive-leaflet-leaf single-node cuttings which were excised fromseven stem positions and grown as single stemmed plants. InRosahybrida ‘Korokis’ Kiss®, ‘Tanettahn’Manhattan Blue®, and ‘Sweet Promise’ Sonia®,following release of the buds from apical dominance by excision,morphogenetic development was studied until anthesis. The timefrom excision/planting until onset of bud growth, visible flowerbud appearance, and anthesis was generally shorter in plantsoriginating from apical bud positions than from basipetal positions.Topophysis mainly affected the onset of axillary bud growth;the earliest growth and development was found in cuttings fromthe second uppermost node position. This node tended to havethe lowest plastochron value, which indicated the existenceof a transition between sylleptic and proleptic buds. Stem lengthat visible flower bud and at anthesis generally increased asthe cutting position changed basipetally until the second lowestposition, and the number of five-leaflet-leaves at anthesisand the total number of nodes generally increased basipetally.For internode length, growth rate, and fresh biomass efficiencythe cuttings taken from the uppermost and lowermost positionsgenerally had significantly lower values than cuttings fromall medial positions. At anthesis, plants originating from cuttingsexcised from lower medial positions generally had a higher freshweight, greater flower stem diameter, and a significantly higherspecific fresh weight than those plants originating from apicalor basal positions. Among the cultivars, Sonia was the mostefficient in increasing fresh biomass and had the highest growthrate, whereas Manhattan Blue possessed the highest specificfresh weight, indicating a higher plant quality. It is suggestedthat topophysis inRosa is an independent phenomenon intrinsicto the axillary bud. apical dominance; axillary bud growth; fresh biomass accumulation; cut rose; flowering; Rosaceae; Rosa hybrida L.; rose; shoot growth; single-stem roses; specific fresh weight; topophysis; quality     

Apical Dominance in the Tomato: Some Further Observations on Isogenic Lines Showing Varying Degrees of Side-shoot Development

A study has been made of the distribution of substances resemblingindol-3-ylacetic acid (IAA), abscisic acid (ABA) and cytokinin-likesubstances in the stem tissue of Craigella tomato plants ascompared with that found in two isogenic lines of this variety,Craigella Blind (blbl) and Craigella Lateral Suppressor (lsls),in both of which side shoot growth is suppressed to varyingdegrees. There was no evidence to suggest that the distributionof these hormones in the stem had any association with the differentpatterns of side shoot development of the three types, thoughsome of the lateral suppressor plants which exhibited only partialbud inhibition did show a relation between high auxin and abscisicacid levels and lack of side shoot development from the centralnodes of the shoot. Decapitation led to a stimulation of bud outgrowth from allnodes of the Craigella plants but the lateral suppressor plantsremained inhibited. The blind plants were found to initiatebud primordia at the cotyledonary nodes only when the severedapex was replaced by exogenous IAA. The results are discussed in relation to our knowledge of themechanisms controlling apical dominance in the tomato. Lycopersicon esculentumL, tomato, apical dominance, growth regulation, indol-3-ylacetic acid, abscisic acid, cytokinins     

拟南芥矮小丛生突变体的分离与分子鉴定

顶端优势是指侧生分生组织的生长被主茎或主花序所抑制.最近的研究通过分离和鉴定顶端优势发生改变的突变体开始揭示顶端优势的分子机制.通过T-DNA标签法分离了拟南芥矮小丛生(bushy and dwarf 1, bud1 )突变体.突变体植株的表型包括顶端优势丧失、株型矮小,表明bud1 突变体存在生长素代谢、运输或信号传导的缺陷.一个对生长素特异反应的启动子驱动的报告基因在bud1 中表达模式改变.生长素敏感性和运输能力的测定表明这两个过程在 bud1中均正常.以上结果显示bud1 表型是生长素代谢缺陷的结果.遗传分析表明BUD1 为半显性突变且与一个T-DNA插入共分离,可通过iPCR方法分离.     

拟南芥矮小丛生突变体的分离与分子鉴定

顶端优势是指侧生分生组织的生长被主茎或主花序所抑制。最近的研究通过分离和鉴定顶端优势发生改变的突变体开始揭示顶端优势的分子机制。通过T-DNA标签法分离了拟南芥矮小丛生(bushy and dwarf l,budl)突变体。突变体植株的表型包括顶端优势丧失、株型矮小,表明budl突变体存在生长素代谢、运输或信号传导的缺陷。一个对生长素特异反应的启动子驱动的报告基因在budl中表达模式改变。生长素敏感性和运输能力的测定表明这两个过程在budl中均正常。以上结果显示budl表型是生长素代谢缺陷的结果。遗传分析表明BUDI为半显性突变且与一个T-DNA插入共分离,可通过iPCR方法分离。     

Gravistimulus Direction, Ethylene Production and Shoot Elongation in the Release of Apical Dominance in Pharbitis nil

Prasad, T. K. and Cline, M. G. 1985. Gravistimulus direction,ethylene production and shoot elongation in the release of apicaldominance in Pharbitis nil.—. exp. Bot. 36: 1969–1975.Release of apical dominance can be induced in Pharbitis nilby the inversion of the upper shoot. This promotion of outgrowthof the highest lateral bud adjacent to the bend of the stemappears to be mediated by ethylene inhibition of growth of theinverted main shoot. In the present investigation the existenceof a direct correlation between ethylene evolution and the directionof gravistimulus is demonstrated as well as an inverse correlationbetween ethylene production by the inverted upper shoot andits elongation. An inverse correlation also exists between elongationof the inverted upper shoot and the outgrowth of the highestlateral bud if the lower portion of the shoot (below the bend)is oriented in an upright position. The latent period for shoot–inversioninduction of ethylene production is about 2 h. These resultssupport the hypothesis of indirect ethylene control of apicaldominance release by retardation of elongation of the invertedshoot. Key words: Shoot inversion, gravistimulus, ethylene, latent period, bud outgrowth, pharbitis nil     

Does Auxin Play a Role in the Release of Apical Dominance by Shoot Inversion in Ipomoea nil?

According to the auxin-inhibition hypothesis of apical dominance,apically produced auxin moves down the stem and inhibits axillarybud outgrowth, either directly or indirectly. This hypothesishas been examined further by monitoring changes in basipetalauxin transport and endogenous auxin concentration in Ipomoeanil caused by shoot inversion, a stimulus that releases apicaldominance. The results indicate that inversion reduces auxintransport in the main stem. In upright shoots of intact plants,a 16-h pretreatment with [³H]IAA 4 cm below the apex resultsin downward movement of label and accumulation in nodes, especiallythe cotyledonary node. Label does not accumulate in the lateralbuds. GC-MS determinations of endogenous free auxin level inthe fourth node, where a lateral bud grows out following inversionof the upper part of the shoot, show no changes at 3 and 8 hafter inversion, the range of times for inversion-induced budrelease, or at 24 h, when bud outgrowth is continuing. However,inversion did cause a just-detectable decrease (approx. 10%)in the IAA level of the shoot's elongation region. Althoughauxin transport in segments of the main stem is partially inhibitedby inversion over a period shorter than the latent time of budrelease, thus providing a means for the expected depletion ofauxin in the fourth node, no depletion could be detected there.These results suggest that either a decrease in IAA level inthe main stem is not causal of bud release or that the decreasedIAA pool responsible for bud release is compartmented and cannotbe measured in whole-tissue extracts.Copyright 1993, 1999 AcademicPress Apical dominance, auxin content, auxin transport, axillary bud release, GC-MS, Ipomoea nil, Pharbitis nil, shoot inversion     

Effects of Abscisic Acid on the Growth Pattern of the Shoot Apical Meristem and on Flowering in Chenopodium rubrum L.

Abscisic acid (ABA) at 1 x 10^–4 M or 3 x 10^–4 Mwas applied to the apical buds of Chenopodium rubrum plantsexposed to different photoperiodic treatments and showing differentpatterns of floral differentiation. Stimulation of growth inwidth of the apical meristem of the shoot and/or inhibitionof growth in length was obtained under all photoperiodic treatments.This change of growth pattern was followed by different effectson flowering. In non-induced plants grown under continuous light ABA stimulatedpericlinal divisions in the peripheral zone and the initiationof leaves as well as the growth in width of bud primordia. Inplants induced by two short days reduced growth of the meristemcoincided with ABA application. Longitudinal growth of the meristemwas inhibited in this case and only a temporary stimulationof inflorescence formation took place. In plants induced ata very early stage, ABA exerted a strong inhibitory effect onflowering. A permanent and reproducible stimulatory effect onflowering was obtained in plants induced by three sub-criticalphotoperiodic cycles if ABA was applied to apices released fromapical dominance. In this case formation of lateral organs andinternodes was promoted by ABA and was followed by stimulatedinflorescence formation. Gibberellic acid (GA2) at 1x 10^–4M or 3 x 10^–4 M brought about a similar effect on floweringas ABA, although the primary growth effect was different, i.e.GA₂ stimulated longitudinal growth. The effects of ABA and GA₂ on floral differentiation have beencompared with earlier results obtained from auxin and kinetinapplications. These growth hormones are believed to regulateflowering by changing cellular growth within the shoot apex.Depending on the actual state of the meristem identical growthresponses may result in different patterns of organogenesisand even in opposite effects on flowering. Shoot apex, flowering, photoperiodic induction, abscisic acid, gibberellic acid, Chenopodium rubrum L.     

Development and Growth Potential of Axillary Buds in Roses as Affected by Bud Age

The effect of axillary bud age on the development and potentialfor growth of the bud into a shoot was studied in roses. Ageof the buds occupying a similar position on the plant variedfrom 'subtending leaf just unfolded' up to 1 year later. Withincreasing age of the axillary bud its dry mass, dry-matterpercentage and number of leaves, including leaf primordia, increased.The apical meristem of the axillary bud remained vegetativeas long as subjected to apical dominance, even for 1 year. The potential for growth of buds was studied either by pruningthe parent shoot above the bud, by grafting the bud or by culturingthe bud in vitro. When the correlative inhibition (i.e. dominationof the apical region over the axillary buds) was released, additionalleaves and eventually a flower formed. The number of additionalleaves decreased with increasing bud age and became more orless constant for axillary buds of shoots beyond the harvestablestage, while the total number of leaves preceding the flowerincreased. An increase in bud age was reflected in a greaternumber of scales, including transitional leaves, and in a greaternumber of non-elongated internodes of the subsequent shoot.Time until bud break slightly decreased with increasing budage; it was long, relatively, for 1 year old buds, when theysprouted attached to the parent shoot. Shoot length, mass andleaf area were not clearly affected by the age of the bud thatdeveloped into the shoot. With increasing bud age the numberof pith cells in the subsequent shoot increased, indicatinga greater potential diameter of the shoot. However, final diameterwas dependent on the assimilate supply after bud break. Axillarybuds obviously need a certain developmental stage to be ableto break. When released from correlative inhibition at an earlierstage, increased leaf initiation occurs before bud break.Copyright1994, 1999 Academic Press Age, axillary bud, cell number, cell size, pith, shoot growth, Rosa hybrida, rose     

Concepts and terminology of apical dominance

Apical dominance is the control exerted by the shoot apex over lateral bud outgrowth. The concepts and terminology associated with apical dominance as used by various plant scientists sometimes differ, which may lead to significant misconceptions. Apical dominance and its release may be divided into four developmental stages: (I) lateral bud formation, (II) imposition of inhibition on lateral bud growth, (III) release of apical dominance following decapitation, and (IV) branch shoot development. Particular emphasis is given to discriminating between Stage III, which is accompanied by initial bud outgrowth during the first few hours of release and may be promoted by cytokinin and inhibited by auxin, and Stage IV, which is accompanied by subsequent bud outgrowth occurring days or weeks after decapitation and which may be promoted by auxin and gibberellin. The importance of not interpreting data measured in Stage IV on the basis of conditions and processes occurring in Stage III is discussed as well as the correlation between degree of branching and endogenous auxin content, branching mutants, the quantification of apical dominance in various species (including Arabidopsis ), and apical control in trees.