Gravistimulus direction, ethylene production and shoot elongation in the release of apical dominance in Pharbitis nil

Release of apical dominance can be induced in Pharbitis nil by the inversion of the upper shoot. This promotion of outgrowth of the highest lateral bud adjacent to the bend of the stem appears to be mediated by ethylene inhibition of growth of the inverted main shoot. In the present investigation the existence of a direct correlation between ethylene evolution and the direction of gravistimulus is demonstrated as well as an inverse correlation between ethylene production by the inverted upper shoot and its elongation. An inverse correlation also exists between elongation of the inverted upper shoot and the outgrowth of the highest lateral bud if the lower portion of the shoot (below the bend) is oriented in an upright position. The patent period for shoot-inversion induction of ethylene production is about 2 h. These results support the hypothesis of indirect ethylene control of apical dominance release by retardation of elongation of the inverted shoot.     

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     

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 Role of Gravity in Apical Dominance : Effects of Clinostating on Shoot Inversion-Induced Ethylene Production, Shoot Elongation, and Lateral Bud Growth

Shoot inversion-induced release of apical dominance in Pharbitis nil is inhibited by rotating the plant at 0.42 revolutions per minute in a vertical plane perpendicular to the axis of rotation of a horizontal clinostat. Clinostating prevented lateral bud outgrowth, apparently by negating the restriction of the shoot elongation via reduction of ethylene production in the inverted shoot. Radial stem expansion was also decreased. Data from experiments with intact tissue and isolated segments indicated that shoot-inversion stimulates ethylene production by increasing the activity of 1-aminocyclopropane-1-carboxylic acid synthase. The results support the hypothesis that shoot inversion-induced release of apical dominance in Pharbitis nil is due to gravity stress and is mediated by ethylene-induced retardation of the elongation of the inverted shoot.     

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     

Shoot inversion-induced ethylene in pharbitis nil induces the release of apical dominance by restricting shoot elongation

Shoot inversion induces outgrowth of the highest lateral bud (HLB) adjacent to the bend in the stem in Pharbitis nil. In order to determine whether or not ethylene produced by shoot inversion plays a direct role in promoting or inhibiting bud outgrowth, comparisons were made of endogenous levels of ethylene in the HLB and HLB node of plants with and without inverted shoots. That no changes were found suggests that the control of apical dominance does not involve the direct action of ethylene. This conclusion is further supported by evidence that the direct application of ethylene inhibitors or ethrel to inactive or induced lateral buds has no significant effect on bud outgrowth. The hypothesis that ethylene evolved during shoot inversion indirectly promotes the outgrowth of the highest lateral bud (HLB) by restricting terminal bud (TB) growth is found to be supported by the following observations: (1) the restriction of TB growth appears to occur before the beginning of HLB outgrowth; (2) the treatment of the inverted portion of the shoot with AgNO₃, an inhibitor of ethylene action, dramatically eliminates both the restriction of TB growth and the promotion of HLB outgrowth which usually accompany shoot inversion; and (3) the treatment of the upper shoot of an upright plant with ethrel mimics shoot inversion by retarding upper shoot growth and inducing outgrowth of the lateral bud basipetal to the treated region.     

Gibberellin-enhanced elongation of inverted Pharbitis nil shoot prevents the release of apical dominance

Ethylene evolution resulting from the gravity stress of shoot inversion appears to induce the release of apical dominance in Pharbitis nil (L.) by inhibiting elongation of the inverted shoot. It has been previously demonstrated that this shoot inversion release of apical dominance can be prevented by promoting elongation in the inverted shoot via interference with ethylene synthesis or action. In the present study it was shown that apical dominance release can also be prevented by promoting elongation of the inverted shoot via treatment with gibberellic acid (GA₃). A synergistic effect was observed when AgNO₃, the ethylene action inhibitor, was applied with GA₃. Both GA₃ and AgNO₃ increased ethylene production in the inverted shoot. These results are consistent with the view that it is ethylene-induced inhibition of elongation and not any direct effect of ethylene per se which is responsible for the outgrowth of the highest lateral bud.     

Shoot inversion-induced ethylene production: a general phenomenon?

Shoot inversion induction of ethylene production was found in inverted shoots of corn, peas, soybean, sunflower, tomato, andPharbitis nil. The increases in ethylene production were found to range from two- to threefold in soybean to eightfold in corn and sunflower. The occurrence of peaks of ethylene production ranged from 16 h following shoot inversion in corn to 72 h inPharbitis. That the enhanced ethylene production was due to activation of 1-aminocyclopropane-1-carboxylic acid (ACC) synthase is supported by the finding of increased ACC content in inverted shoots of all species tested. Shoot inversion inhibition of elongation was found in inverted shoots of pea, soybean, sunflower, tomato, andPharbitis nil. This inhibition is thought to be mediated via increased ethylene production in the inverted shoots. That shoot inversion induction of ethylene is not a persistent effect is supported by the finding that ethylene synthesis could be terminated by reorientation of shoots to the upright position and could be reinitiated by the subsequent inversion of the shoots. The effects of shoot inversion on the enhancement of ethylene production and on the inhibition of elongation of the inverted shoot appear to be general phenomena.     

Timing of Growth Inhibition following Shoot Inversion in Pharbitis nil

Shoot inversion in Pharbitis nil results in the enhancement of ethylene production and in the inhibition of elongation in the growth zone of the inverted shoot. The initial increase in ethylene production previously was detected within 2 to 2.75 hours after inversion. In the present study, the initial inhibition of shoot elongation was detected within 1.5 to 4 hours with a weighted mean of 2.4 hours. Ethylene treatment of upright shoots inhibited elongation in 1.5 hours. A cause and effect relationship between shoot inversion-enhanced ethylene production and inhibition of elongation cannot be excluded.     

Shoot Inversion Inhibition of Stem Elongation in Pharbitis nil: A Possible Role for Ethylene-Induced Glycoprotein and Lignin

Inversion of the upper shoot of Pharbitis nil results in the inhibition of elongation in the inverted stem. The objective of the present study was to determine how shoot inversion-induced gravity stress inhibited elongation and to elucidate the possible role of ethylene-induced glycoprotein and lignin in this process. Determinations of hydroxyproline, peroxidase, phenylalanine ammonia-lyase (PAL), phenol, and lignin content/activity were carried out by appropriate spectrophotometric methods. It was found that inversion and Ethrel treatments of upright shoots caused significant increases in hydroxyproline content, peroxidase, and PAL activity in 12 hours and in phenol and lignin contents in 24 hours. All of these increases except for that of cytoplasmic peroxidase activity were partially reversed by AgNO₃, the ethylene action inhibitor. It is concluded that possible cross-linking associated with the accumulation of ethylene-induced hydroxyproline-rich glycoprotein and lignin may be responsible for the later stages of cessation of elongation in the inverted Pharbitis shoot.     

Shoot Inversion-induced Ethylene Production in the Diageotropica Tomato Mutant

Shoot inversion was found to inhibit shoot elongation over 24h in the diageotropica (dgt) mutant tomato and its isogenicparent, VNF8, by 55% and 51%, respectively. Previous studieswith normal tomato and other species would suggest that gravitystress-induced ethylene production in the inverted shoot retardselongation. Since exposure of the diageotropic shoot of thedgt mutant tomato to ethylene restores normal upward growth,it might appear that dgt is defective in its capacity to produceethylene. That shoot inversion did stimulate some ethylene productionand that the ethylene action inhibitor, AgNO₃, largely negatedthe inhibiting effect of shoot inversion on elongation in dgtstrongly suggest that the ethylene produced in the invertedshoot was responsible for its retardation of growth. Althoughethylene production in the slower growing dgt was much lessthan that in the faster-growing VNF8, the dgt shoot was foundto be much more sensitive to ethylene. Shoot inversion, ethylene, shoot elongation, diageotropica (dgt), auxin     

Floral initiation under continuous light in Pharbitis nil, a typical short-day plant

Seedling-cuttings of Pharbitis nil, a typical short-day plant,initiated floral buds under continuous light of 2200–2400lux at 24–26?C. When cultured under poor-nutritional conditions,the node bearing the first floral bud was as low as the 4thone. A close relation between floral initiation under continuouslight and retarded vegetative growth was observed. (Received September 28, 1973; )     

Flowering Response in Relation to C and N Contents of Pharbitis nil Plants Cultured in Nitrogen-Poor Media

Flowering of seedlings of Pharbitis nil, strains Violet andTendan, cultured in modified White's medium, was promoted bymedium dilution, the critical dark period being shortened byabout 15 min. Dilution of the N source alone was enough to causethe medium-dilution effect. Dilution of the culture medium duringthe day before and on the day of exposure to the dark-period(a total of two days) caused the maximum dilution effect. TheC and N contents of the cotyledons and of the shoot apices changedrapidly in response to medium dilution. In 1/2-strength White'smedium with 1/1,000 strength NO₃^– which was most effectivefor flower promotion, the C-N ratio was highest. In 1/2-strengthmodified White's medium, in which flowering was lowest withthe longest critical dark period, the C-N ratio was lowest.Thus, there is a close relation between flowering response andthe C-N ratio in cotyledons or shoot apices of Pharbitis nil. (Received September 14, 1984; Accepted January 26, 1985)     

Author index volume 38 (1985)

Mechanical perturbation (MP, rubbing) of internodes of Pharbitis nil shoots initiates release of lateral buds (LB) from apical dominance within 48 h. Evidence is presented which suggests that MP promotion of LB outgrowth is mediated by ethylene-induced restriction of main shoot growth. Ethylene production in the internodes is stimulated by MP within 2 h. Effects of MP are mimicked by treatments with 1-aminocyclopropane-1-carboxylic acid (ACC) and are negated by the inhibitors of ethylene production or action, aminoethoxy vinylglycine (AVG) and AgNO₃. The fact that effects of MP, ACC and ethylene inhibitors are observed to occur on main shoot growth at least 24 h before they are observed to occur on LB growth suggests a possible cause and effect relationship. MP also causes an increase in internode diameter. MP stimulation of ethylene production appears to be mediated by ACC synthase. The results of this study and our previous studies suggest that apical dominance may be released by any mechanism which induces ethylene restriction of main shoot growth.     

Control of Lateral Bud Growth inPhaseolus vulgarisL. by Ethylene in the Apical Shoot

Elongation growth of the first trifoliate leaf axillary budwas induced by physical restriction of the apical growth orby treating the apical shoot with ethylene or ethephon. Thereis evidence to suggest that the promotion of axillary bud developmentby ethylene action on the apical shoot was associated with theavailability of freely diffusible ethylene in the tissues ofthe treated shoot. Loss of apical dominance was not, apparently,directly dependent on either the internal ethylene concentration(i.e. concentration in the vacuum – extracted gases) oron the rate of ethylene emanation. The effects of aminoethoxyvinylglycine (AVG) and silver nitrate on ethylene production wereexamined. Although treatment of the shoot with tri – iodobenzoicacid (TIBA) induced various morphological responses in the plant,including axillary bud outgrowth, which appeared similar tothe responses to ethylene, the initial effect of TIBA is notthought to be ethylene – mediated.     

Lateral Bud Growth in Phaseolus vulgaris L. and the Levels of Ethylene in the Bud and Adjacent Tissue

Ethephon and the ethylene inhibitors Ag⁺ and aminoethoxyvinylglycine (AVG) inhibited outgrowth of the axillary bud of thefirst trifoliate leaf in decapitated plants of Phaseolus vulgaris.Endogenous ethylene levels decreased in the stem upon decapitationalthough it is not conclusive that a causal relationship existsbetween this decrease and the release of axillary buds frominhibition. The proposition that auxin-induced ethylene is responsiblefor the suppression of axillary bud growth in the decapitatedplant when the apical shoot is replaced by auxin is not borneout in this study. Application of IAA directly to the axillarybud of intact plants gave rise to a transient increase in budgrowth. This growth increment was annulled when AVG was suppliedwith IAA to the bud despite the fact that the dosage of AVGused did not affect the normal slow growth rate of the bud ofthe intact plant or bud outgrowth resulting from shoot decapitation.     

Exogenous Auxin Effects on Lateral Bud Outgrowth in Decapitated Shoots

In 1933 Thimann and Skoog demonstrated exogenous auxin repressionof lateral bud outgrowth in decapitated shoots ofVicia faba. This evidence has given strong support for a role of auxinin apical dominance. Most, but not all, investigators have confirmedThimann and Skoog's results. In the present study, auxin treatmentswere carried out on ten different species or plant types, manyof which were treated with auxin in different forms, media andunder different light conditions. The Thimann–Skoog experimentdid work for most species (i.e. exogenous auxin did repressbud outgrowth) including thedgt tomato mutant which is knownto be insensitive to auxin in certain responses. Toxic auxinsymptoms were observed in some but not all species. The Thimann–Skoogexperiment did not work for greenhouse-grownColeus or forArabidopsis. Light was shown to reduce apical dominance inColeus andIpomoeanil . apical dominance; lateral bud outgrowth; axillary bud; auxin; IAA; decapitation; Vicia faba ; Ipomoea nil ; Pisum sativum ; Phaseolus vulgaris ; Lycopersion exculentum ; dgt ; Coleus blumei ; Arabidopsis thaliana ; Helianthus annuus ; Thimann–Skoog     

Floral initiation of Pharbitis nil, a short-day plant, under continuous high-intensity light

Pharbitis nil, a short-day plant, initiated floral buds undercontinuous illumination at 23°C, provided that the lightintensity was kept at 16,000 lux or above. Stem elongation ofthe plants was strongly inhibited but leaves developed normallyunder this condition. (Received November 26, 1971; )     

Grafting Experiments on Correlative Effects between Lateral Buds

Shoots of Hygrophila sp., which are decussate and have budsof unequal size at a node, were grown in liquid culture. Inexcised nodes it is known that the larger (+) bud inhibits thesmaller (–) bud in the axil of the opposite leaf, andonly one shoot grows out; in nodes split longitudinally bothbuds grow out. When nodes were split and grafted together again(+/– grafts), in general only one bud grew out; if aluminiumfoil was introduced at the nodal region both buds grew out.Thus the inhibitory effect of a + on a – bud is laterallytransmissible across a graft union. In +/– grafts of half-nodesdiffering in age by two plastochrones, a higher proportion yieldedtwo shoots, suggesting that the age differential had some importance.This view is supported by observations on sectioned material.Grafts having two + or two – buds (+/+ grafts) were madebetween half-nodes differing in age by two plastochrones; inthe majority both buds grew out. Thus a + bud inhibits a –bud but usually not another + bud; in either case a considerabledifference in stage of development of the half-nodes may affectthe results. It is concluded that bud dominance resembles apicaldominance, and is probably mediated by hormonal means.     

Inhibition of photoperiodic floral induction in Pharbitis nil by ethylene

Application of ethylene at 100 ppm or higher completely inhibitedflowering in Pharbitis nil when made during an inductive darkperiod. Exposing plants to ethylene before or after the inductivedark period produced only slight or almost no inhibition. Ethylenewas effective when it was applied only to a cotyledon, but wasineffective when applied only to a receptor bud. Ethylene hadno effect on translocation of the floral stimulus. Ethylene-treatedcotyledon did not transfer any flower inhibiting entity. Thus,ethylene is considered to inhibit the induction process(es)in cotyledons. Except for an initial temporary cotyledon epinasty, ethylenetreatment had no effect on the subsequent growth and vigor ofplants. This temporary cotyledon epinasty disappeared withinthe next 24 hr. (Received May 4, 1972; )