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)     

Execution of the auxin replacement apical dominance experiment in temperate woody species

The classic Thimann-Skoog or auxin replacement apical dominance test of exogenous auxin repression of lateral bud outgrowth was successfully executed in both seedlings and older trees of white ash, green ash, and red oak under the following conditions: (1) decapitation of a twig apex and auxin replacement were carried out during spring flush, (2) the decapitation was in the previous season's overwintered wood, and (3) the point of decapitation was below the upper large irrepressible lateral buds but above the lower repressible lateral buds. Although it has been suggested that neither auxin, the terminal bud, nor apical dominance have control over the outgrowth of the irrepressible buds during spring flush, there is evidence in the present study that indicates that such control over the repressible buds exists. In seedlings, second-order branching, which resulted from decapitation of elongating current shoots, was also inhibited by exogenous auxin in the three species. Hence, the auxin replacement experiments did work on year-old proleptic buds (of branches of older trees) that would have entered the bud bank and also on current buds of seedlings. Cytokinin treatments were ineffectual in promoting bud growth.     

Abscisic Acid and Apical Dominance in Phaseolus coccineus L. The Role of Tissue Age

Abscisic acid (ABA) applied to the decapitated second internodeof runner bean plants enhanced outgrowth of lateral buds onlywhen internode stumps were no longer elongating. Applied toelongating internodes of slightly younger plants, ABA causesinhibition of bud outgrowth. Together with 10^–4 M indol-3-ylacetic acid (IAA), ABA stimulated internode elongation and interactedadditively in the inhibition of bud outgrowth. A mixture of10^–5 M ABA and 10^–6 M gibberellic acid (GA₃ ) causedsimilar effects on internode growth as IAA + ABA, but was mutuallyantagonistic in effect on growth of the lateral buds. Abscisic acid, apical dominance, gibberellic acid, indol-3yl acetic acid, Phaseolus coccineus, bean     

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.     

Apical Dominance Control in Ipomoea nil: The Influence of the Shoot Apex, Leaves and Stem

The outgrowth of lateral buds is known to be controlled by theupper shoot tissues, which include the apex, the young leavesand the upper stem. An analysis of the influence of these plantparts on axillary bud elongation in Ipomoea nil was carriedout by various treatments on these specific tissues. A restriction of elongation in the main shoot due to eitherdecapitation or shoot inversion resulted in the release of apicaldominance A non-linear type of compensating growth relationshipwas observed between the 13 cm apical growing region of thestem and the lateral buds. It was determined by decapitation,defoliation and AgNO₃ treatments that both the 13 cm stem-growthregion and the young leaves (1–5 cm in length) had a muchgreater inhibitory influence on the outgrowth of specified lateralbuds than did the stem apex (consisting of the terminal 0.5cm of the shoot). The specified lateral buds which were analyzedfor outgrowth were located a number of nodes below the shootapex. The intervening nodes were debudded. Although the importanceof young leaves in the control of apical dominance has beenpreviously recognized, the most significant result from thepresent study with Ipomoea was the strong influence of the 13cm apical growth region of the stem on the out growth of thelateral buds. Apical dominance, Ipomoea nil L., Pharbitis nil, growth region, lateral bud outgrowth, decapitation, defoliation, shoot inversion     

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.     

Effects of Gibberellin on Shoot Development in the dgt Mutant of Tomato

The morphological effects of gibberellin A₃ (GA₃) on the dgtmutant of tomato were investigated. The mutant effectively showedthe normal range of responses, including a promotion of stemlength due to an increased number of longer internodes, a dramaticincrease in apical dominance, and effects on leaf shape andcolour. In the case of stem elongation, the quantitative responseof the mutant was greater than normal. The morphological abnormalitiescharacteristic of the dgt mutant, such as horizontal growth,a thin stem and hyponastic leaves, were not normalized by GA₃. It is concluded that the demonstrated lack of response to auxinof the dgt mutant does not impair its gibberellin responses. Tomato, gibberellin, auxin, mutant, shoot development     

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     

Effect of gravity on apical dominance in Pharbitis nil.

When the upper part of main shoot of morning glory (Pharbitis nil) is gently bent down, lateral bud on the bending region is released from apical dominance and starts to elongate. But, clinorotating the bending shoots prevents the release of the lateral bud from apical dominance. These results suggest that gravity affects apical dominance in morning glory. Here we verified the gravity-regulated apical dominance by using a weeping morning glory defective in gravitropic response due to abnormal differentiation of endodermis. That is, bending main shoot of the weeping morning glory hardly caused the lateral bud to elongate. In addition, decapitation of apical bud released the lateral bud from apical dominance, and exogenous auxin applied to the cut surface of the decapitated stem was inhibitory to the outgrowth of the lateral bud in the wild type. However, the effect of auxin was much less in the weeping morning glory. Thus, apical dominance of the weeping morning glory was weaker and less influenced by gravity than that of the wild type, which could occur due to abnormal differentiation of endodermis required for graviperception.     

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     

Regulation of Branching in Decussate Species with Unequal Lateral Buds

In the decussate plants Alternanthera philoxeroides and Hygrophilasp. the opposite axillary bud primordia are of unequal sizefrom the time of their inception; the larger or + buds lie alongone helix and the smaller or – buds along another (helicoidalsystem). In decapitated plants of Alternanthera both buds grewout, but unequally; if the node was vertically split growthof the two shoots was more equal, and if the + buds were excisedgrowth of the – shoots approximately equalled that ofcontrol + shoots. In decapitated shoots of Hygrophila grownin sterile culture only one bud, the + or larger one, grew outat each of the upper nodes. In excised cultured nodes, also,only the + bud grew out; but if the nodes were split longitudinallyboth buds grew out, initially rather unequally. These experimentssupport the view that the regulation of branching in these specieshas two components, apical dominance and the dominance of thelarger (+) bud over the smaller (–) bud at the same node.The restriction of growth potentiality imposed on the –bud is not permanent but can be modified. Further correlativeeffects on bud outgrowth include those of the subtending leavesand of buds at other nodes.     

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     

In Vitro Culture and Plant Regeneration in Vicia faba subsp. Equina (var. Spring Blaze)

Explants of stem, leaves, roots, and cotyledons from etiolatedaxenically grown Vicia faba seedlings were cultured on a rangeof media. Shoot organogenesis was only obtained with nodal stemand cotyledonary node explants when cultured on MS medium with3% sucrose, 2.0 mg 1^–1 BAP and 02 mg 1^–1 NAA. Callusproliferation accompanied shoot organogenesis from nodal stemexplants. Successive subculture of nodal stem callus resultedin proliferation of regenerative callus which contained severalshoot bud initials. The capacity for shoot regeneration fromthis callus was maintained for 9 months. Histological studiesreveal de novo formation of meristematic centres in callus andtheir further development into bud primordia. High frequencyrooting of these adventitious shoots was obtained on half-strengthMS medium with 1.5% sucrose, 0.1 mg 1^–1 NAA and 0.5 mg1^–1 kinetin. Key words: Vicia faba, adventitious shoots, axillary shoots, de novomeristem formation, organogenesis, tissue culture     

Apical dominance

Apical dominance is the control exerted by the apical portions of the shoot over the outgrowth of the lateral buds. The classical explanations for correlative inhibition have focused on hormone/nutrient hypotheses. The remarkable progress that has been made in the technology of endogenous hormone quantification in plant tissue has not been accompanied by comparable progress in the elucidation of mechanisms of hormone action in apical dominance. Evidence from hormonal studies suggests that apically produced auxin indirectly suppresses axillary bud outgrowth that is promoted by cytokinin originating from roots/shoots. Significant involvement with other hormones, although less likely, has not been ruled out. Possible changes in tissue sensitivity to hormones should not be overlooked. Auxin-induced oligosaccharide signals originating from the cell walls of shoot tips or polyamines may function as secondary inhibitors to bud growth. Alternatively, apically produced auxin may suppress lateral bud growth by inhibiting auxin export from these buds. Support for a critical role for nutrients in apical dominance keeps resurfacing, especially for auxin-directed nutrient transport and for water as a possible inducing signal for bud outgrowth. Histological and biochemical analyses of lateral buds recently released from apical dominance are urgently needed. The feasibility of manipulating endogenous auxin/cytokinin content in plant tissue by gene insertion and modulation opens the door to exciting approaches as does the use of hormone insensitive/resistant mutants. There is also need to recognize the existence of variability of apical dominance mechanisms among different plant types. The aesthetic and economic implications of understanding apical dominance for the modification of plant structure and form are extremely significant.     

Hormonal control of the outgrowth of axillary buds in <Emphasis Type="Italic">Alstroemeria</Emphasis> cultured <Emphasis Type="Italic">in vitro</Emphasis>

We study apical dominance in Alstroemeria, a plant with an architecture very different from the model species used in research on apical dominance. The standard explant was a rhizome with a tip and two vertically growing shoots from which the larger part had been excised leaving ca. 1 cm stem. The axillary buds that resumed growth were located at this 1-cm stem just above the rhizome. They were released by removal of the rhizome tip and the shoot tips. Replacement of excised tips by lanolin with indole-3-butyric acid (IBA) restored apical dominance. The auxin transport inhibitors 2,3,5-triiodobenzoic acid (TIBA) and N-1-napthylphthalamic acid (NPA) reduced apical dominance. 6-Benzylaminopurine (BAP) enhanced axillary bud outgrowth but the highest concentrations (> 9 μM) caused fasciation. Thidiazuron (TDZ) did not show improvement relative to BAP. Even though the architecture of Alstroemeria and the model species are very different, their hormonal mechanisms in apical dominance are for the greater part very similar.     

Some properties of the amine oxidase in Vicia faba seedlings

Growth of the Vicia faba seedling is accompanied by a rapid15-day increase in amine oxidase activity of the apical parts.Cotyledons and roots were found to be devoid of activity. Thepartially purified enzyme from leaves readily oxidized putrescine,cadaverine, agmatine and spermidine, while dopamine (3-hydroxytyramine)and L- and D-lysine were oxidized more slowly. The Km valueswere 1.9?10^–3 M for cadaverine, 3.7?10^–5 M for putrescine,7.8?10^–4 M for spermidine, and 5.9?10^–3 M for dopamine.Carbonyl reagents and copper-binding agents were effective inhibitorsof Vicia faba amine oxidase. The diethyldithiocarbamate-treatedenzyme could be reactivated specifically by cupric copper. (Received May 25, 1977; )     

Studies on the Geotropism of Roots: III. EFFECTS OF GEOTROPIC STIMULATION ON GROWTH-SUBSTANCE CONCENTRATIONS IN VICIA FABA ROOT TIPS

The auxins contained in 5-mm. tips of horizontal Vicia fabaroots have been compared with those in tips of vertical rootsafter cold ethanol extraction, paper-chromatographic separation,and Avena mesocotyl bioassay. At about the time curvature commencesin horizontal roots there is a marked increase in the contentof an auxin corresponding to ‘AP(ii)’ of pea roots(Rf 0.35–0.65 in isobutanol/methanol/water). There areindications that this is not due to its release from an inactivebound state but that it is either synthesized de novo or maybe converted from another auxin corresponding to ‘AP(iii)’of pea roots (Rf 0.75–1.0). The literature dealing with the auxins of geotropically stimulatedorgans is reassessed and it is concluded that, with the exceptionof the Avena coleoptile, there is very little evidence favouringa simple transport redistribution of auxin under gravity; themajority of the data favour an effect of gravity on auxin metabolism.     

A Comparative Auxin and Cytokinin Study in Normal and to-2 Mutant Tomato Plants

Torosa-2 (to-2), a tomato mutant with strong apical dominance,was studied in order to determine the mechanism of shoot outgrowthcontrol. In decapitated or defoliated to-2 plants only a fewshoots grew and IAA or morphactin application had little oronly short term effects. No differences were found in auxinand cytokinin activities between normal and mutant plants upto 20 days after sowing. In the period from 40 to 90 days theIAA content increased equally in both genotypes. During thesame period, however, cytokinin increased only in normal plants.The results obtained with the to-2 mutant indicate that budsdo not shoot, probably because bud differentiation does notoccur. The ratio of auxin to cytokinin clearly was affectedby the low level of cytokinin in all tissues of the to-2 plants.This led us to the conclusion that insufficient quantities ofcytokinin for lateral bud differentiation is the cause of thestrong apical dominance in to-2. (Received January 20, 1982; Accepted April 26, 1982)     

A Microautoradiographic Study of Auxin Transport in the Stem of Intact Pea Seedlings (Pisum sativum L.)

Soluble-compound microautoradiography was used to determinethe distribution of radioactivity in transverse sections ofintact dwarf pea stems (Pisum sativum L.) following the applicationof [³H]IAA to the apical bud. Near the transport front labelwas confined to the cambial zone of the axial bundles, includingthe differentiating secondary vascular elements. Fully differentiatedphloem and xylem elements remained unlabelled and no radioactivitywas detected in the leaf or stipule traces. Similar resultswere obtained in experiments with Vicia faba L. plants. Nearerthe labelled apical bud of the pea there was a more generaldistribution of label and evidence was found of free-space transportof radioactive material in the pith. When [³H]IAA was applied to mature foliage leaves the greatestconcentration of label was found in the differentiated phloemelements of the appropriate leaf trace and in the phloem ofthe adjacent axial bundles. Both basipetal and acropetal transportwas detected in this case. These results are consistent with the conclusions drawn fromearlier transport experiments which indicated that in the intactplant the long-distance basipetal transport of auxin from theapical bud takes place in a system which is separated from thephloem transport system and suggests that the vascular cambiumand its immediate derivatives may function as the normal pathwayfor the longdistance movement of auxin in the plant. The physiologicalsignificance of such a transport system for auxin is discussed.