Effect of apex excision and replacement by 1-naphthylacetic acid on cytokinin concentration and apical dominance in pea plants

As known from literature lateral buds from pea ( Pisum sativum ) plants are released from apical dominance when repeatedly treated with exogenous cytokinins. Little is known, however, about the endogenous role of cytokinins in this process and whether they interact with basipolar transported IAA, generally regarded as the main signal controlling apical dominance. This paper presents evidence that such an interaction exists.
The excision of the apex of pea plants resulted in the release of inhibited lateral buds from apical dominance (AD). This could be entirely prevented by applying 1-naphthylacetic acid (NAA) to the cut end of the shoot. Removal of the apex also resulted in a rapid and rather large increase in the endogenous concentrations of zeatin riboside (ZR), isopentenyladenosine (iAdo) and an as yet unidentified polar zeatin derivative in the node and internode below the point of decapitation. This accumulation of ZR and iAdo, was strongly reduced by the application of NAA. The observed increase in cytokinin concentration preceded the elongation of the lateral buds, suggesting that endogenous cytokinins play a significant role in the release of lateral buds from AD. However, the effect of NAA on the concentration of cytokinins clearly demonstrated the dominant role of the polar basipetally transported auxin in AD. The results suggest a mutual interaction between the basipolar IAA transport system and cytokinins obviously produced in the roots and transported via the xylem into the stem of the pea plants.     

Stimulatory effect of cytokinins and interaction with IAA on the release of lateral buds of pea plants from apical dominance

Lateral buds of pea plants can be released from apical dominance and even be transformed into dominant shoots when repeatedly treated with synthetic exogenous cytokinins (CKs). The mechanism of the effect of CKs, however, is not clear. The results in this work showed that the stimulatory effects of CKs on the growth of lateral buds and the increase in their fresh weights in pea plants depended on the structure and concentration of the CKs used. The effect of N-(2-chloro-4-pyridyl)-N'-phenylurea (CPPU) was stronger than that of 6-benzylaminopurine (6-BA). Indoleacetic acid (IAA) concentration in shoot, IAA export out of the treated apex and basipetal transport in stems were markedly increased after the application of CPPU or 6-BA to the apex or the second node of pea plant. This increase was positively correlated with the increased concentration of the applied CKs. These results suggest that the increased IAA synthesis and export induced by CKs application might be responsible for the growth of lateral shoots in intact pea plants.     

Involvement of auxin and CKs in boron deficiency induced changes in apical dominance of pea plants (Pisum sativum L.)

It has previously been shown that boron (B) deficiency inhibits growth of the plant apex, which consequently results in a relatively weak apical dominance, and a subsequent sprouting of lateral buds. Auxin and cytokinins (CKs) are the two most important phytohormones involved in the regulation of apical dominance. In this study, the possible involvement of these two hormones in B-deficiency-induced changes in apical dominance was investigated by applying B or the synthetic CK CPPU to the shoot apex of pea plants grown in nutrient solution without B supply. Export of IAA out of the shoot apex, as well as the level of IAA, Z/ZR and isopentenyl-adenine/isopentenyl-adenosine (i-Ade/i-Ado) in the shoot apex were assayed. In addition, polar IAA transport capacity was measured in two internodes of different ages using 3H-IAA. In B-deficient plants, both the level of auxin and CKs were reduced, and the export of auxin from the shoot apex was considerably decreased relative to plants well supplied with B. Application of B to the shoot apex restored the endogenous Z/ZR and IAA level to control levels and increased the export of IAA from the shoot apex, as well as the 3H-IAA transport capacity in the newly developed internodes. Further, B application to the shoot apex inhibited lateral bud growth and stimulated lateral root formation, presumably by stimulated polar IAA transport. Applying CPPU to the shoot apex, a treatment that stimulates IAA export under adequate B supply, considerably reduced the endogenous Z/ZR concentration in the shoot apex, but had no stimulatory effect on IAA concentration and transport in B-deficient plants. A similar situation appeared to exist in lateral buds of B-deficient plants as, in contrast to plants well supplied with B, application of CKs to these plants did not stimulate lateral bud growth. In contrast to the changes of Z/ZR levels in the shoot apex, which occurred after application of B or CPPU, the levels of i-Ade/i-Ado stayed more or less constant. These results suggest that there is a complex interaction between B supply and plant hormones, with a B-deficiency-induced inhibition of IAA export from the shoot apex as one of the earliest measurable events.     

Enhancement of lateral bud growth in Coleus blumei BENTH. by (2-chloroethyl) trimethylammonium chloride (CCC)

The relationship of GA to apical dominance in Coleus was examinedby substituting 1 % IAA, in lanolin, for the shoot apex of CCC-treated,control and GA-treated plants containing, theoretically, hyponormal,normal and hypernormal GA levels, respectively. The greatestinhibition of lateral bud growth was obtained in the treatmentcombining 1 % IAA and 100 ppm GA, suggesting that GA may beimportant in the apical dominance of Coleus. CCC inhibited main axis growth, reduced the level of endogenousGA and caused a marked release of lateral buds from apical dominance. The significant stimulation of lateral bud growth by CCC couldnot be ascribed to reduced endogenous GA since it was not reversedby exogenous GA, or by GA plus IAA, whereas 100 ppm GA overcamethe inhibition of main axis growth by CCC. It was also shownthat the CCC stimulation was not a result of compensatory growth,that is, enhanced lateral bud growth resulting from reducedapical bud growth. The CCC effect on lateral buds was interpretedas involving a system independent of auxin and GA or else apossible immobilization of auxin in addition to inhibition ofGA biosynthesis. (Received December 5, 1967; )     

Changes in abscisic acid and indole-3-acetic acid in axillary buds of Elytrigia repens released from apical dominance

Growth of axillary buds on the rhizomes of Elytrigia repens (L) Nevski is strongly dominated by the rhizome apex, by mechanisms which may involve endogenous hormones. We determined the distribution of indole-3-acetic acid (IAA) and abscisic acid (ABA) in rhizomes and measured (by gas-chromatography-mass spectrometry) their content in axillary buds after rhizomes were decapitated. The same measurements were also made in buds induced to sprout by removing their subtending scale leaves. The ABA content tended to be higher in the apical bud and in the axillary buds than in the adjacent internodes, and tended to decline basipetally in the internodes and scale leaves. IAA was similary distributed, except that there was less difference between the buds and other rhizome parts. After rhizomes were decapitated, the ABA content of the first axillary bud declined to 20% of that of control values within 24 h, while the IAA content showed no marked tendency to change. The ABA content also declined within 12 h in the first axillary bud after rhizomes were denuded, while the content of IAA tended to increase after 6 h. These changes occurred before the length of the first axillary bud increased 24–48 h after rhizomes were decapitated or denuded. We conclude that the release of axillary buds from apical dominance in E. repens does not require IAA content to be reduced, but is associated with reduced ABA content.     

Changes of endogenous hormones in lateral buds of chrysanthemum during their outgrowth

The character of branching for two chrysanthemum (Chrysanthemum × morifolium) cvs. Jinghai and Jingyun was observed, and the changes of endogenous hormones in apical and lateral buds were investigated to determine the relationship between the pattern of hormone distribution, apical dominance, and lateral bud outgrowth. The growth rate of Jinghai lateral buds was higher than that of Jingyun. In vegetative growth stage, IAA level in apical buds of Jingyun was significantly higher than in Jinghai. After flower induction, IAA level in apical buds of two cultivars decreased remarkably, but the IAA level decreased in Jingyun faster than in Jinghai. These results showed that the higher was the IAA level in apical buds the stronger was inhibition of lateral bud outgrowth. An increase in IAA and iP/iPA and a decrease in ABA concentrations were closely associated with lateral bud growth alterations in chrysanthemum.     

Changes after Decapitation in Concentrations of Indole-3-Acetic Acid and Abscisic Acid in the Larger Axillary Bud of Phaseolus vulgaris L. cv Tender Green

Early changes in the concentrations of indole-3-acetic acid (IAA) and abscisic acid (ABA) were investigated in the larger axillary bud of 2-week-old Phaseolus vulgaris L. cv Tender Green seedlings after removal of the dominant apical bud. Concentrations of these two hormones were measured at 4, 6, 8, 12 and 24 hours following decapitation of the apical bud and its subtending shoot. Quantitations were accomplished using either gas chromatography-mass spectrometry-selected ion monitoring (GS-MS-SIM) with [¹³C₆]-IAA or [²H₆]-ABA as quantitative internal standards, or by an indirect enzyme-linked immunosorbent assay, validated by GC-MS-SIM. Within 4 hours after decapitation the IAA concentration in the axillary bud had increased fivefold, remaining relatively constant thereafter. The concentration of ABA in axillary buds of decapitated plants was 30 to 70% lower than for buds of intact plants from 4 to 24 hours following decapitation. Fresh weight of buds on decapitated plants had increased by 8 hours after decapitation and this increase was even more prominent by 24 hours. Anatomical assessment of the larger axillary buds at 0, 8, and 24 hours following decapitation showed that most of the growth was due to cell expansion, especially in the intermodal region. Thus, IAA concentration in the axillary bud increases appreciably within a very few hours of decapitation. Coincidental with the rise in IAA concentration is a modest, but significant reduction in ABA concentration in these axillary buds after decapitation.     

STUDIES ON THE FLORAL HISTOGENESIS AND PHYSIOLOGY OF PERILLA. III. EFFECTS OF INDOLEACETIC ACID ON THE FLOWERING OF APICAL BUDS AND EXPLANTS IN CULTURE

Raghavan , V. (Princeton U., Princeton, N. J.) Studies on the floral histogenesis and physiology of Perilla. III. Effects of indoleacetic acid on the flowering of apical buds and explants in culture. Amer. Jour. Bot. 48(10): 870–876. Illus. 1961.—The responses of apical buds and explants of a short-day plant, Perilla frutescens (L.) Britt. var. 'Tall Late,' when grown in vitro in White's medium supplemented with indoleacetic acid (IAA) and subjected to short-days (SD) or long-days (LD), are described. Additions of varying concentrations of IAA to the medium inhibited the flowering of the photoinduced apical buds in 2 ways. There was a progressive delay in the appearance of the first signs at the apex and a gradual transition from the more flower-like structures in lower concentrations of IAA (0.1 mg/liter) to sterile cones in higher doses. The sterile cones had florets with well-developed calyx and corolla lobes, but lacked the sporogenous tissues. When subjected to LD, visible signs were observed only in buds grown in 0.1 and 1.0 mg/liter IAA, the pronounced effect of the auxin being in the step-wise inhibition in the formation of the non-sporogenous tissues of the differentiating florets. Flowering of the explants with the 1st pair of unfolded leaves was also inhibited by IAA in either SD or LD, but the 1st signs appeared relatively faster than in apical buds. When photoinduced, explants with the 1st and 2nd pairs of unfolded leaves flowered in all concentrations of IAA tried (up to 100 mg/liter) while those kept in LD remained entirely vegetative.     

Branch development in Lupinus angustifolius L.#II. Relationship with endogenous ABA, IAA and cytokinins in axillary and main stem buds

The concentrations of indole-3-acetic acid (IAA), cytokinins (CK) and abscisic acid (ABA) were measured in buds of different regions (main stem and lateral branches) of Lupinus angustifolius L. (cv. Merrit) and at different stages in the development of branches. In lupin, branching patterns are the result of discrete regions of axillary branches (upper, middle and basal) which elongate at much different rates. Early in development only the main shoot elongates, followed usually by basal branch growth and then rapid upper branch growth. Branches in the middle of the main stem grow only weakly or fail to develop. Levels of IAA were generally high in the apical buds of slowly growing branches and low in buds from strongly growing branches, whereas CK levels showed the opposite relationship. CK:IAA ratio showed a closer relationship with the rate of growth of a particular branch better than the levels of either CK or IAA alone. During early stages of growth ABA concentration did not follow the rate of branch growth. However, later in development, where growth did not closely match the ratio of CK:IAA, ABA level showed a strong negative relationship with growth. A significant decrease in ABA was associated with continued strong growth of the main stem apex following a decline in CK:IAA ratio. Overall, the best relationship between the level of growth factors in apical buds and branching pattern in lupin was the ratio of CK:IAA, implying that high CK:IAA at a given bud would promote growth. ABA level appeared to play a secondary role, as a growth inhibitor.     

Levels of endogenous indole-3-acetic acid and indole-3-acetylaspartic acid during adventitious root formation in pea cuttings

Levels of endogenous indole-3-acetic acid (IAA) and indole-3-acetylaspartic acid (IAAsp) were monitored in various parts of leafy cuttings of pea ( Pisum sativum L. cv. Marma) during the course of adventitious root formation. IAA and IAAsp were identified by combined gas chromatography—mass spectrometry, and the quantitations were performed by means of high performance liquid chromatography with spectrofluorometric detection. IAA levels in the root forming tissue of the stem base, the upper part of the stem base (where no roots were formed), and the shoot apex remained constant during the period studied and were similar to levels occurring in the intact seedling. A reduction of the IAA level in the root regenerating zone, achieved by removing the shoot apex, resulted in almost complete inhibition of root formation. The IAAsp level in the shoot apex also remained constant, whereas in the stem base it increased 6-fold during the first 3 days. These results show that root initiation may occur without increased IAA levels in the root regenerating zone. It is concluded that the steady-state concentration is maintained by basipetal IAA transport from the shoot apex and by conjugation of excessive IAA with aspartic acid, thereby preventing accumulation of IAA in the tissue.     

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     

The effect of leaf primordia and auxin on leaf initiation rate in strawberry

Summary Removal of the leaf primordia hastens the rate of leaf initiation at the apex, and a paste containing 0.2% IAA in lanoline will substitute for the effect of the leaf primordia. Physical factors involved in the alteration in bud structure resulting from defoliation, such as gaseous diffusion and shading from light, have only negligible effect on the rate of leaf initiation, and the compsition of the internal atmosphere of the intact buds is not very different from the external atmosphere.The evidence suggests that developing leaf primordia inhibit cell division of the apical meristem through their production of auxin which is discharged into the stem at points which are morphologically basal to the apical cells; this, therefore, could be another case of correlative inhibition by auxin, comparable with the inhibition of lateral buds by the terminal apex.     

Changes in endogenous hormones in apple during bud burst induced by defoliation

Under the tropical conditions of East Java, terminal buds of apple burst at any time of the year in response to removal of the subtending leaves. Following two such defoliations, two weeks apart on separate trees, there was a decrease in abscisic acid (ABA), a three-fold increase in gibberellin-like substances (GAs) and only a slight increase in cytokinin-like substances (CKs) in the apex tissue of closed buds. These changes preceded bud opening and the associated increases in fresh and dry weight, and may be causally related to bud burst. In open buds (i.e. young expanding leaves) the concentration of CKs was greater, and the concentrations of ABA and GAs less, than the concentrations in closed buds. As the leaves expanded, ABA increased and GAs and CKs decreased in concentration. The decrease in concentration of GAs and CKs, however, was due to the rise in dry weight of the expanding tissue; the amounts of all three hormones (per apex) increased. During bud burst there was a concurrent decrease in the CKs of subtending stems, suggesting a transfer into the expanding bud tissues. Removal of the old leaves by defoliation may remove the source of ABA and allow the amount of GAs in the apex to rise, bud burst following. Stem CKs may be utilized in the expansion of the new leaves in the bursting buds.     

Effect of Morphactin, Gibberellic Acid and Auxin and on Growth and Development of Bryophyllum tubiflorum

six month old LD- and SD- grown plants of Bryophyllum tubiflorum were treated with morphactin (n-buty1-9-hydroxy-fluorene-(9)-carboxylate), singly and in combination with GA₃ and IAA. Morphactin and IAA decreased stem elongation and number of leaves, the effect increasing with concentration concentation. Morphactin also caused lateral buds to develop into branches and the fusion of upper leaves to form structures of different shapes. GA₃ enhanced stem elongation, increased leaf number and induced floral buds under SD conditions. It reversed the inhibitory effect of morphactin on stem elongation, leaf number and leaf fusion and also restored apical dominance when applied simultaneously with morphactin. The stimulaneous application of IAA also reversed the morphactin effects on leaf fusion and on apical dominance. The results have been discussed in the light of literature available on the subject.     

Factors Influencing the Distribution of Growth Between Stem and Axillary Buds in Decapitated Bean Plants

Phaseolus multiflorus plants at three stages of developmentwere decapitated either immediately below the apical bud orlower down at a point 1 cm above the insertion of the primaryleaves. Growth regulators in lanolin were applied to the cutstem surface. IAA always inhibited axillary bud elongation anddry-matter accumulation, and enhanced internode dry weight butnot elongation. GA₃ applied below the apical bud greatly increasedinternode elongation and dry weight, but simultaneously reducedbud elongation and dry-weight increase. Application of GA₃ 1cm above the buds had no effect on bud elongation in the youngestplants, but enhanced their elongation in the two older groups.IAA always antagonized GA₃-enhancement of internode extensiongrowth, whereas its effects on GA₃-enhanced dry-matter accumulationdepended on the stage of internode development. Bud elongationwas greater in plants treated with GA₃+IAA than in plants treatedonly with IAA, except in the youngest plants decapitated immediatelybelow the apical bud, where GA₃ caused a slight increase inIAA-induced bud inhibition. GA₃ increased inhibition of buddry weight by IAA in the two youngest groups of plants, butslightly reduced it in the oldest plants. No simple compensatorygrowth relationship existed between internode and buds. It wasconcluded that, (1) auxin appears to be the principal growthhormone concerned in correlative inhibition, and (2) availabilityof gibberellin to internode and buds is of importance as a modifyingfactor in auxin-regulated apical dominance by virtue of itslocal effects on growth in the internode and in the buds.     

Effects of boron starvation on boron compartmentation, and possibly hormone-mediated elongation growth and apical dominance of pea (Pisum sativum) plants

Two experiments were carried out to study the effects of boron (B) deficiency on 7-day-old pea plants for 6 or 9 days under controlled growth chamber conditions. Growth and apical dominance (AD) of the plants and their B concentration and compartmentation were followed throughout the starvation period. Additionally, auxin (indoleacetic acid, IAA) concentration in the shoot apex and polar transport from it were measured along with the cytokinin (CK) concentration in the shoot apex and the roots. The results demonstrate that during a 6-day B-deficiency period, B concentration in the water-insoluble residue of the roots was very stable and could not easily be reduced. In contrast, B concentration in the cell sap fraction was very sensitive to external B supply. Twelve hours after transferring the plants from B-sufficient to B-deficient solutions, the B concentration in root cell sap declined to half the concentration of the control plants. In addition, B concentration in the new aerial plant parts, which developed after the onset of the B-deficiency treatment, was extremely low. A decline in elongation growth could be observed as soon as about 4 days after the imposition of B deficiency. This preceded the first measurable growth of lateral buds (release from AD). Before the onset of these morphological changes, there was a considerable decline in CK concentration, accompanied by a dramatic decrease in IAA export out of the shoot apex, a decline in IAA concentration in the shoot apex and the roots and a reduced capacity for polar IAA-transport. These changes are discussed as possible reasons for the observed reduction in elongation growth and AD. These hormonal changes themselves are possibly the result of the decreased symplasmic B concentration, which in turn may be responsible for the reduced concentration in apical CKs. A sequence of events, which may be causally related, is suggested to explain the effects of B deficiency on the growth and AD of pea plants.     

香荚兰花芽分化至萌发期内源激素的变化

以香荚兰 (Vanillafragrans)为材料 ,研究不同栽培条件下花芽分化和萌发期内源激素变化 ,分析和探讨内源激素在花芽分化和萌发中的作用 ,香荚兰花芽分化时期茎里的激素含量降低 ,芽里激素含量升高 ,其中相对高的ZR和ZR ABA有利于分化 ,IAA和IAA ABA的一定增加也利于分化 ,过高或没有IAA的增加则不利于花芽分化。大多数花芽形成于倒垂茎蔓上 ,花芽分化期 (11～ 12月 ) ,倒垂茎蔓的茎里生长类激素含量降低大于竖立茎蔓 ,芽的激素含量增高则多于竖立茎蔓 ,倒垂茎蔓的这种变化可能是有利于花芽分化。香荚兰生长中顶端优势明显 ,去顶后侧芽里ZR、GA、IAA增高 ,这与 11～ 12月去顶促进倒垂茎蔓开花可能有关。     

The Mechanism of Apical Dominance in Coleus

The development of lateral buds in isolated stems of Coleus blumei is inhibited by low concentrations of indoleacetic acid or other auxins, just as in other plants. The inhibition can be fully reversed by kinetin, about 3 times as much kinetin as IAA being needed. However, the outgrowth of the same lateral buds on intact Coleus plants is sensitive to environmental conditions, well-nourished plants in full daylight often showing little inhibition by applied auxin. It is shown that (a) the solvent used for IAA, (b) the light intensity and (c) the nitrogen and phosphorus nutrition, all control the sensitivity of the buds to auxin inhibition. Using water instead of lanolin, lowering the light intensity or decreasing the supply of either nitrogen or phosphorus all increase the degree of apical dominance.     

Reversal of auxin induced inhibition by ethrel

The interaction between exogenous 2-chloroethylphosphonic acid (Ethrel, CEPA) and auxin (both native and synthetic—IAA) was studied on pea and bean seedlings, potato tubers, and processed flax plants. After the addition of ethrel the inhibiting effect of IAA was decreased in all objects and it was found that the concentration of the growth of the regulators played an important role. The growth response of a part of flax hypocotyl, as induced by exogenous auxin produced in the cotyledon, was reversed by ethrel, too. The application of ethrel on the epicotyl apex in beans resulted in the lost of apical dominance of epicotyl and in the growth of lateral buds together with the epicotyl. When stimulating the growth, ethrel reverses the inhibitions through the decrease in the auxin content (from an inhibiting, supraoptimum level to an optimum one which already stimulates growth). In objects with a low content of endogenous auxin the ethrel induced the decrease in the auxin content and shows an inhibiting effect on growth.