The Effect of (2-chloroethyl)-trimethylammonium Chloride on the Growth of Barley

Barley (Hordeum vulgare L. C.I.666) was shown to be susceptibleto the growth retardant (2-chloroethyl)-trimethylammonium chloride(CCC). The estimation of cell number in the dwarfed third leafblade indicated that a decrease in mitotic activity had occurredin treated plants. There was also a decrease in cell size intreated plants. The dwarfing action of CCC was reversed by exogenousgibberellic acid (GA₃) but this was shown to be the result ofincreased cell elongation only. GA₃ did not promote cell divisionin healthy or CCC-treated plants. Assay of endogenous gibberellinsshowed a significant reduction in the level of a substance correspondingto GA₃ in CCC-treated plants. It is suggested that CCC-induceddwarfing of barley is largely the result of a reduction in meristematicactivity. This may be related to an effect on gibberellin biosynthesisbut is not reversed by the application of exogenous GA ₃.     

Growth Hormones and the Creeping Thistle Rust

A re-examination of the life cycle of the rust fungus Puccimapunctiformis (Str.) Rohl. (syn. P. suaveolens (Pers.) Rostr.,P. obtegens Tul. ex Fuckel) on the creeping thistle Cirsiumarvense (L) Scop, has led to the development of a method forgrowing comparable healthy and diseased thistle plants in thegreenhouse. These were used to define the developmental changeswhich result from systemic infection. Closely similar changesare produced by the exogenous application of gibberellic acid(GA₃). The endogenous gibberellin content of systemically diseasedthistle plants is consistently higher than that of healthy plantsduring the earlier part of the season when they are growingfaster. Evidence is given for the presence of two gibberellinswhose relative proportions change during development. The firsthas not been characterized but there is some evidence that thesecond may be GA₃. These results support the hypothesis thatsome of the characteristic features of systemic infection ofthe host are due to the excess production of gibberellins.     

Mechanical Stress and Gibberellin: Regulation of Hollowing Induction in the Stem of a Bean Plant, Phaseolus vulgaris L.

In pole bean plants, mechanical stress (MS) inhibited stem elongationand induced radial thickening of the stem. Application of uniconazole,an inhibitor of gibberellin biosynthesis, also reduced stemgrowth but had no effect on stem diameter. Both MS and uniconazolesignificantly reduced hollowing of the first internodes, butonly the former increased ethylene evolution from the firstinternode. Application of GA₃ increased the length of the firstinternode and decreased its diameter in bush bean plants; thiswas accompanied by a significant promotion of stem hollowing.Aminooxyacetic acid (AOA) decreased ethylene evolution fromthe GA₃-treated internodes, though it did not reduce the GA₃-inducedhollowing of the first internodes. Application of GA₃ affectedneither ethylene evolution nor cellulase activity in the firstinternodes of bush bean plants. Application of GA₃ stimulatedmuch greater cell elongation in the center of pith tissue thanin the outer surrounding tissues, suggesting a possible physicalbreakage of the inner cells, which leads the hollowing of beanstems. These results suggest that gibberellin is a factor responsiblefor stem hollowing in bean plants. Because MS is known to reducegibberellin content in bean plants [Suge (1978) Plant Cell Physiol.21: 303] MS may inhibit stem hollowing by reducing the amountof endogenous gibberellin. (Received July 1, 1994; Accepted November 8, 1994)     

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.     

Internode Length in Pisum. A New Allele at the le Locus

In Pisum sativum L. a third, more severe, allele at the internodelength locus le is identified and named le^d. Plants homozygousfor le^d possess shorter internodes and appear relatively lessresponsive to GA₂₀ than comparable le (dwarf) plants. Gene le^dmay act by reducing the 3ß-hydroxylation of GA₂₀ tothe highly active GA₁ more effectively than does gene le. Theresults indicate that le is a leaky mutant and therefore thatendogenous GA₁ influences internode elongation in dwarf (le)plants. Pisum sativum, peas, internode length, genetics, gibberellin, dwarf elongation     

Dwarfism Caused by Floral-Bud Removal in Petunia and its Reversal by Gibberellin

The effect of floral-bud removal at different stages of developmenton the plant height and on the total number of buds of Petuniawas studied. Continuous removal of all the floral buds 2 d beforeanthesis caused a marked decrease in plant height and also increasedthe total number of floral buds formed thereafter. At otherstages of floral bud development, bud removal had a lesser effecton both phenomena. Moreover, the plants did not respond to budremoval at anthesis. GA₃ at 25 ppm applied to plants from which the buds had beenremoved, promoted stem elongation. The most pronounced effectwas on plants from which the buds were removed 2 d before anthesis,but it had no effect on plants from which the buds were removedat anthesis stage. The possible involvement of endogenous growth hormones in theresponse of Petunia plants to floral-bud removal and to applicationof GA₃ is discussed. Bud removal, bud number, dwarfness, GA₃, Petunia, plant height     

The effects of temperature and the Rht3 dwarfing gene on growth, cell extension, and gibberellin content and responsiveness in the wheat leaf

The effects of low temperature and the Rht3 dwarfing gene onthe dynamics of cell extension in leaf 2 of wheat were examinedin relation to gibberellin (GA) content and GA-responsivenessof the extension zone. Leaf 2 of wild-type (rht3) wheat closelyresembled that of the Rht3 dwarf mutant when seedlings weregrown at 10C. The maximum relative elemental growth rate (REGR)within the extension zone in both genotypes was lower at 10Cthan at 20C, but the position with respect to the leaf basewas unaffected by temperature. The size of the extension zoneand epidermal cell lengths were similar in both genotypes at10C. Growth at 20C, instead of 10C, increased the lengthof the extension zone beyond the point of maximum REGR in thewild type, but not in the Rht3 mutant. Increasing temperatureresulted in longer epidermal cells in the wild type. Treatingwild-type plants at 10C with gibberellic acid (GA₃) also increasedthe length of the extension zone, but the Rht3 mutant was GA-non-responsive.However, the concentrations of endogenous GA₁ and GA₃ remainedsimilar across the extension zone of wild-type plants grownat both temperatures, despite large differences in leaf growthrates. The period of accelerating REGR as cells enter the extensionzone, and the maximum REGR attained, are apparently not affectedby GA. It is proposed that GA functions as a stimulus for continuedcell extension by preventing cell maturation in the region beyondmaximum REGR and that low temperature increases the sensitivitythreshold for GA action. Key words: Cell extension, gibberellin, Rht3 dwarfing gene, temperature, wheat leaf     

Gibberellin A12 Is Converted to Gibberellin A53 in Cultured Cells of Nicotiana tabacum and Catharanthus roseus

The metabolism of GA₁₂ and its precursors was investigated incultured cells of seven cell lines of Nicotiana tabacum andthree cell lines of Catharanthus roseus using ^l4C-labeled substrates.The presence of a metabolic pathway from ent-7-hydroxykaurenoicacid to GA₅₃ via GA₁₂-aldehyde and GA₁₂ was demonstrated inthe cultured cells. GA₁₂ was effectively converted to GA₅₃ incells of BY-2, 2b-4, 2b-13 and CG from N. tabacum. By contrast,GA₅₃ was not converted to any other GAs in all of the linesof cells examined. The metabolism of C₁₉-GAs was also examinedusing ³H-labeled substrates. The conversion of GA₂₀ to GA₂₉and GA, and of GA₄ to GA₃₄ occurred more efficiently in cellsfrom C. roseus than in cells from N. tabacum. However, 13-hydroxylationof GA₄ and GA₉ was not observed in any of the cell culturesexamined. Among the various metabolites, GA₅₃, GA₂₉ and GA₃₄were identified by full-scan GC/MS. (Received December 20, 1990; Accepted May 27, 1991)     

Long-Day Induced Bud Break in Salix pentandra Is Associated with Transiently Elevated Levels of GA1 and Gradual Increase in Indole-3-Acetic Acid

Initiation of dormancy in woody species is induced by a shortphotoperiod. In the early stages of dormancy development, growthcan be initiated by long days. To study the possible involvementof gibberellins and indole-3-acetic acid in bud break in Salixpentandra, effects on levels of these plant hormones of transferof seedlings in an early stage of dormancy to a growth-inductivelong photoperiod was investigated. The bud break and initiationof growth correlated with a rapid transient increase in GA₁in defined zones of shoot tips to levels twice of that in continuouslyelongating long day-grown control seedlings, followed by a rapiddecrease to levels similar to in these control plants. Also,the contents of GA₅₃, GA₁₉, GA₂₀, GA₈ increased upon transferfrom short to long days. Levels of indole-3-acetic acid showeda gradual decline under short days, and increased graduallyupon transfer to long days up to a level of continuously elongatingplants. The present data suggest an interaction between gibberellinand indole-3-acetic acid in the photoperiodic control of dormancydevelopment and bud break in S. pentandra. (Received October 14, 1996; Accepted February 18, 1997)     

Gibberellin-Induced Flowering in Cordyline (Agavaceae)

Normal, terminal inflorescences of Cordyline terminalis, a woodymonocotyledon, appeared 4–6 weeks after apical buds weretreated with gibberellin A₃ (GA₃) or GA₄₊₇. There was no responseto GA₁₃. Large plants, newly rooted cuttings, and seedlingsall responded, although there were clonal differences. Floweringwas induced under natural day lengths throughout the year. Untreatedcontrol plants in all experiments remained vegetative. Dracaenaspp. did not respond to the three gibberellins.     

The Anther Smut of Sea Campion: A Study of the Role of Growth Regulators in the Dwarfing Symptom

The sympton of dwarfing in the sea campion, Silene vulgaris(Moench) Garcke sub sp. maritima(With.) A. & D. Löveinfected with the anther smut fungus Ustilago violacea (Pers.)Fuckel, a systemic, perennial parasite, has been investigated. Extracts of both healthy and diseased plants contain IAA andGA₃, but diseased plants contain less gibberellins than healthyplants. Neither IAA nor gibberellins were detected in significantquantities in the medium when U. violacea was grown in pureculture, but IAN was present. IAN was also found m extractsof diseased plants and it is tentatively suggested that it isformed by the fungus and may accumulate in the host owing tothe inability of the plant to convert IAN to IAA The exogenousapplication of IAN to healthy plants does not produce any diseasesymptoms. The dwarfing symptom of the diseased plant may be due to thelower levels of gibberellins which it contains compared withhealthy plants since the exogenous application of GA₃, restoresthe diseased plant to normal growth     

Growth Regulators and the Effect of Barley Yellow Dwarf Virus on Barley (Hordeum vulgare L.)

Estimation of cell number in the third leaf of barley (Hordeumvulgare L. C I 666) infected with barley yellow dwarf virus(BYDV) showed a marked decrease in the mitotic activity of theinfected plants Assay of endogenous gibberellins revealed adecrease in the level of a substance corresponding to gibberellicacid (GA₃) in BYDV-infected plants No significant differencein the level of endogenous auxins was observed Application ofgibberellic acid to infected plants reversed the dwarfing effectbut the response was not significantly different from the responseof healthy plants and was found to be due to increased cellelongation. It is suggested that the dwarfing of BYDV-infectedplants is a result of reduced mitotic activity This may be relatedto the reduced level of endogenous gibberellins.     

Evidence that Daminozide, but not Two Other Growth Retardants, Modifies the Fate of Applied Gibberellin A9 in Chrysanthemum morifolium Ramat

Interactions between three growth retardants and gibberellinA₉ (GA₉) in Chrysanthemum morifolium Ramat. cv. Bright GoldenAnne grown in short days have revealed that the particular retardantemployed modifies the response to this gibberellin. When plants were given a foliar spray of daminozide (a substitutedsuccinamic acid), the effects of a subsequent application ofGA₉ to each lateral shoot, on both stem length and the rateof inflorescence development, were relatively small. However,GA₉ was highly active on plants previously sprayed with piproctanylbromide (a quaternary ammonium, piperidinium compound) or onthose given a compost drench of chlorphonium chloride (a quaternaryphosphonium retardant). In the absence of GA₉ all three retardantsreduced stem length and delayed flowering. Irrespective of the extent of the responses to various dosesof GA₉ (1–50 µg per shoot), a close relationshipwas maintained between internode extension and earliness offlowering, as in previous studies with retardants and othergibberellins. On plants which had received chlorphonium chlorideGA₃ and GA₁₃ were highly active and poorly active, respectively. The data suggest that daminozide could act as a retardant notonly by blocking the biosynthesis of gibberellins but also,wholly or partly, by restricting the metabolism or action ofendogenous gibberellins.     

Cell Number and Cell Size in Parthenocarpicvs. Pollinated Blueberry (Vaccinium ashei) Fruits

Gibberellic acid (GA₃) promotes parthenocarpic fruit developmentand is used commercially to increase fruit set in many crops.However, fruit size is usually smaller than that of pollinatedfruit. The purpose of this work was to determine the anatomicalbasis for differences in fruit size between pollinated and GA₃-inducedparthenocarpic blueberry (Vaccinium asheiReade) fruits. Freshweights at ripening averaged 1.6 and 2.5 g for GA₃-treatedvs.pollinated fruits, respectively. In both pollinated and GA₃-treatedfruits, mesocarp cell number comprised about 75% of the totalpericarp cell number, and increased from     

Mitotic Activities and Levels of Nuclear DNA in the Apical Meristem of Silene armeria (strain SI.2) Following Application of Gibberellin A3

Strain S1.2 of Silene armeria was grown under an 8h-photoperiodand treated with GA₃ every day for 20 days. This growth substancecaused stem elongation, but no flowering in this long-day plant.Changes in the mitotic index and DNA content of cells in thevarious zones of the apical meristem before, during and afterGA₃ treatment were described. Mitotic activity and increasein the proportion of nuclei at the 4C level (S+G₂ phase of thecell cycle) were strongly stimulated in the rib-meristem, andto a lesser extent in the lateral zone, but not in the axialzone. This stimulation of apical activity reached a peak aftertwo GA₃ treatments and then declined gradually, so that after20 days the activity in GA₃-treated meristems was lower thanthat in untreated controls; at this point most cells were inthe G₁ phase. When the GA₃ treatment was discontinued, there was a gradualincrease in the mitotic activity which ultimately reached thesame level as that in controls. Stem elongation ceased and leavesformed aerial rosettes. It is concluded that in vegetative plants of strain S1.2 ofSilene armeria GA₃ acts mainly on the rib-meristem cells whichresults in stem elongation. Lack of response in the axial cellsexplains why GA₃ fails to induce flowering in this strain ofSilene armeria. (Received June 18, 1983; Accepted August 3, 1983)     

Unusual Branching in the Seedlings of Lotus japonicus--Gibberellins Reveal the Nitrogen-sensitive Cell Divisions within the Pericycle on Roots

We studied the effects of several plant-growth regulators onthe induction of nodule-like structures on roots of Lotus japonicus,which has been proposed as a candidate for a leguminous plantfor molecular genetic analysis. Contrary to our expectations,the addition of gibberellin A₃ (GA₃) at concentrations of 10^-4M to 10^-4 M resulted in the formation of nodule-like structureson roots when seedlings were plated on nitrogen-free Fahraeusagar medium. GA₄ also induced such outgrowths but was less activethan GA₃. Application of an inhibitor of auxin transport, N-(1-naphthyl)-phthalamicacid (NPA) and of kinetin, which have been reported to inducepseudonodules in other legumes, had no effect on L. japonicus.Microscopic observations showed that GA₃-induced nodule-likestructures were caused by cell divisions within the pericycleon the roots. In addition, the outgrowths elicited by GA₃ couldbe completely suppressed by the addition of 15 mM potassiumnitrate or ammonium nitrate. These results show that the pericyclecells of the roots of L. japonicus are specifically sensitiveto gibberellins and that potential for cell division might bemodulated by nitrogen compounds. We also examined the effectsof ancymidol and uniconazole [S-3307; (E)-1-(4-chIorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol],two synthetic plant-growth retardants. Both compounds at 3 x10^-5 M significantly increased the number of stunted lateralroots. The unusual branching could not be counteracted by theexogenous addition of GA₃ but by 10^-6 M brassinolide. We discussthe physiological role of brassinolide in the initiation oflateral roots. (Received August 4, 1995; Accepted March 11, 1996)     

Physiological Controls on the Production of Cleistogamous and Chasmogamous Flowers in Lamium amplexicaule L

Plants of Lamium amplexicaule, grown under short-day field conditionsin Northern California produce predominantly closed flowers.Under long-day field conditions, plants may produce up to 50per cent open flowers, the same total number of flowers beingproduced under each daylength regime. A 20 ml drop of GA₃ or GA₇ (100µM) was applied to themain shoot apex of plants growing under short-day conditions,and all subsequently produced flowers opened. CCC, an inhibitorof gibberellin synthesis, was applied (0.06 per cent) to thesoil of seedlings grown under long-day conditions. The CCC-treatedplants were dwarfed and bore only closed flowers. With GA₇ appliedexogenously to the CCC-treated shoots, inhibition was released,resulting in elongated internodes and open flower production.The timing of flower production and internodal extension inLamium amplexicaule are positively correlated. When floral primordiawere removed from main shoots, the average internode lengthsdecreased. The number of nodes produced in treated plants wasincreased as a result of flower removal. Exogenous GA₇ (10 µM)applied to the nodes from which flowers were removed resultedin internodal extension but had no effect on node number. Twoprocesses that may contribute to the control of the productionof open and closed flowers in Lamium amplexicaule are: (1) anincreasing anther sac size from lower to upper node flowersthat may exert control locally, via GN production, on corollaexpansion, and (2) a photoperiodic control. Lamium amplexicaule L, henbit, cleistogamy, chasmogamy, gibberellins, gibberellic acid (GA₃), (2 chloroethyl) trimethylammonium chloride (cycocel)     

A Crucial Role for Gibberellins in Stress Protection of Plants

The hypothesis that there is an intimate relationship betweengibberellin levels and plant stress protection has been testedusing near-isogenic lines of a normal and dwarf barley (Hordeumvulgare L.). Application of paclobutrazol (a triazole), inducedstress protection in the normal line and application of GA₃to the dwarf reversed the inherent stress tolerance. Reversalof the dwarf phenotype by specific gibberellins (GAs) suggeststhat the conversion of GA₂₀ to GA₁ and GA₉ to GA₄ has been compromised.These observations indicate that modulation of specific GAsplays a key role in stress protection. The preferential useof non-chemical technologies including enzyme regulation, phytochromeA overexpression, and breeding for induction of stress tolerancein plants are discussed. (Received November 26, 1998; Accepted March 11, 1999)     

Effect of gibberellic acid on starch content of soybean (Glycine max L.) and its correlation with extension growth

This investigation was undertaken to study the effect of GA₃on extension growth of Glycine max L. and on starch contentof its individual internodes at maturity. The effect on hydrolyticactivity of the extract of different internodes was also studied.GA₃ stimulates the extension growth of stem by increasing theelongation of those internodes which are either in the processof elongation or being differentiated at the time of treatment.Starch content decreases with the position of the internode(from base upwards) on the intact plant. Corresponding internodeshave minimum starch content in 100 ppm GA₃-treated plants andmaximum in the controls. Internodes which show the maximum elongationdue to GA₃ treatment, show the least starch content and alsoshow maximum hydrolytic activity during the period of elongation.It is suggested that enhanced extension growth is brought aboutby enhanced mobilization of reserve food by GA₃. (Received November 21, 1967; )     

IAA-oxidase in Impatiens balsamina Affected by GA3 and Tannic Acid

The activity of IAA-oxidase increased in the leaves of Impatiensbalsamina plants receiving inductive photoperiodic cycles andin plants receiving treatments with gibberellic acid (GA₃) and/ortannic acid (TA), even under non-inductive photoperiods; theactivity also increased in the stem receiving inductive photoperiodiccycles (8 h). Treatment with GA₃ and TA mimics the effect ofSD cycles in the development of some isoenzymes of IAA-oxidase.Thus a new isoenzyme at R_f 0.48 developed in the leaves andone at R_f 0.82 developed in both the stem and the leaves ofall plants receiving inductive treatments – photoperiodicor chemical – but not in water-treated controls undernon-inductive photoperiods. Another isoenzyme at R_f 0.68 developedonly in the stems. Flowering, gibberellic acid, IAA oxidase, Impatiens, phenols, photoperiod