The effect of plant growth regulator treatments on the levels of ethylene emanating from excised turgid and wilted wheat leaves

Abscisic acid (ABA) inhibits the production of ethylene induced by water stress in excised wheat leaves and counteracts the stimulatory effect of 6-benzyladenine (BA) on this process. The stimulatory effect of BA and the inhibitory effect of ABA were equally pronounced whether external or endogenous ethylene levels were determined. When leaves were sprayed or floated on solutions of BA, indole-3-acetic acid (IAA), gibberellic acid (GA₃), or ABA, the relative activities of these growth regulators on stress-induced ethylene at 10^-4 mol l^-1 were BA>IAA >GA₃>controls>ABA. In non-stressed leaves, however, where the levels of ethylene produced were 2–20 times smaller, the relative activities were IAA >BA>GA₃>controls>ABA. The effects of BA and ABA spray treatment on water stress induced ethylene were closely similar whether the solutions were applied 2 or 18 h prior to the initiation of water stress. The relationships between the levels of endogenous growth regulators in the plant and ethylene release induced by water stress are discussed.Abbreviations BA 6-benzyladenine - IAA indole-3-acetic acid - GA₃ gibberellic acid - ABA abscisic acid - GLC gas-liquid chromatography - _leaf leaf water potential     

Evidence for the accumulation of a germination inhibitor during progressive thermoinhibition of seeds of celery (Apium graveolens L.)

The germination of seeds of celery (Apium graveolens L.) becomes progressively thermoinhibited on incubation in the dark at high temperatures, the inhibitory temperature being dependent on the cultivar used. In two high-dormancy cultivars of celery, the production of germination inhibitors in seeds incubated in the dark at 26°C gradually increased over a 7-day period. Inhibitor production was measured by incubating seeds of the low-dormancy cultivar Florida 683 in homogenates of the thermoinhibited seeds of the high-dormancy cultivars and recording germination either in the light or with the gibberellins A₄ and A₇ (GA_4/7) in the dark. Most Florida 683 seeds which failed to germinate in the homogenates after 15 days were induced to germinate by addition of N⁶-benzyladenine (BA). The presence of BA in addition to GA_4/7 throughout incubation in the dark completely overcame the inhibitory effects of homogenates. This indicates that thermoinhibition of celery seeds is associated with the accumulation of a germination inhibitor which interacts with cytokinins. This does not appear to be abscisic acid (ABA) since ABA levels in thermoinhibited seeds were lower than in untreated seeds and did not increase with duration of high temperature treatment.Abbreviations ABA Abscisic acid - BA N⁶-benzyladenine - GA_4/7 a mixture of the gibberellins A₄ and A₇ - HTP high-temperature pretreatment     

Role of acid efflux during growth promotion of primary leaves of Phaseolus vulgaris L. by hormones and light

The white-light-(WL) induced enlargement of dicotyledonous leaf cells is known to occur via an acid-growth mechanism; i.e., WL causes leaf cells to excrete protons which lead to an increase in wall extensibility and thus cell enlargement. Gibberellic acid (GA₃) and N⁶-benzyladenine (BA) also induce leaf cell enlargement. To see if they also act via acid-induced cell wall loosening, a comparison has been made of WL-, GA₃-and BA-induced growth of strips, taken from primary leaves of bean (Phaseolus vulgaris L.) plants raised in continuous red light for 10 d. White light, GA₃ and BA all increased wall extensibility as measured by the Instron technique, and this change preceded the increase in growth rate. However, whereas WL induced significant proton excretion, neither GA₃ nor BA caused any acidification of the apoplast. Furthermore, neutral buffers, which effectively inhibited the growth induced by WL, were without effect on growth promoted by either GA₃ or BA. These results indicate that while WL, GA₃ and BA all initiate growth in bean leaves by altering cell-wall properties, GA₃ and BA do so through some wall loosening mechanism other than wall acidification. Neither gibberellin nor cytokinin is likely to play a major role in light-induced cell enlargement of dicotyledonous leaves.Abbreviations BA N^o-benzyladenine - FC fusicoccin - GA₃ gibberellic acid - RL red light - SK medium 10 mM sucrose+10mM KCl - WL white light     

The effect of 6-benzyladenine and leaf ageing treatment on the levels of stress-induced ethylene emanating from wilted wheat leaves

Leaf ageing was initiated in wheat leaves by floating excised leaves on distilled water in darkness for up to three days. After a given period of time the leaves were blotted and wilted to a leaf water potential (_leaf) of approximately — 12 · 10² k Pa and then placed in a sealed chamber so that the stress-induced ethylene could be determined. The longer the period of leaf ageing the smaller were the levels of stress-induced ethylene. Treatments which are known to delay leaf senescence, such as floating the leaves on solutions of 6-benzyladenine (BA) or in the light instead of in the dark, were found to partly restore or even enhance the stress-induced ethylene levels. For example leaves allowed to age for 1 day whilst floating on 10^-4 mol l^-1 BA solution produced up to 200% more ethylene than freshly harvested leaves when both were subjected to water stress treatment. When BA solutions were applied as foliar sprays to wheat seedlings, the day before wilting treatment, the amount of stress-induced ethylene diffusing from the freshly harvested leaves (i.e. no leaf ageing treatment) was significantly increased. This occurred when water stress was induced in the leaves by either immersing the roots of seedlings in carbowax solution or by wilting excised leaves in a stream of warm air. There was a substantial synergistic effect between BA treatment and water stress in relation to the amount of ethylene diffusing from the leaves. This effect was 7.5-fold in experiments where excised leaves were floated on 10^-4 moll^-1 BA solutions for 1 day prior to wilting and 4.7-fold for intact plants where the BA solution was applied as a foliar spray the day before water stress induction by carbowax. We can postulate from the experiments in this paper that leaf ageing (and/or the depletion of an ethylene substrate, probably a photosynthetic product or a substance derived from it) and the level of endogenous cytokinin are probably important factors which determine the amount of ethylene emanating from leaves during water stress. Moreover, the results suggest that wheat shoots may contain sub-optimal levels of cytokinins in regard to their potential ability to produce ethylene under stress.Abbreviations BA 6-benzyladenine - IAA indole-3-acetic acid - IAAsp indole-3-acetyl-N-aspartate - GLC gas-liquid chromatography - _leaf leaf water potential     

Implication of Ethylene in the Release of Secondary Dormancy in Amaranthus caudatus L. Seeds by Gibberellins or Cytokinin

Ethephon (Eth), gibberellin A₃, A_{4 + 7} (GA₃, GA_{4 + 7}), and 6-benzyladenine (BA) removed secondary dormancy of Amaranthus caudatus seeds. The GAs and BA potentiated the effect of ethephon or 1-aminocyclopropane-1-carboxylic acid (ACC), an ethylene biosynthesis precursor, in terms of the rate or final percent of germination. Aminoethoxyvinylglycine (AVG), an ACC synthase activity inhibitor, was observed to simultaneously inhibit the release from dormancy effected by GA₃ or BA as well as the ethylene production stimulated by these regulators. Breaking of secondary dormancy by GA₃, GA_{4 + 7} or BA was prevented by 2,5-norbornadiene (NBD), an inhibitor of ethylene binding. Ethylene completely or markedly reversed the inhibitory effect of NBD. We thus conclude that the removal of secondary dormancy in Amaranthus caudatus seeds by gibberellin or benzyladenine involves ethylene biosynthesis and action.     

Dormancy break of celery (Apium graveolens L.) seeds by plant derived smoke extract

Seed dormancy of a highly-dormant cultivar of celery (Apium graveolens L.) was broken by combinations of plant-derived smoke extract or N⁶-benzyladenine (BA) and gibberellins A_4/7 (GA_4/7) in the dark at temperatures between 18 and 26°C. A less dormant cultivar which responded to GA_4/7 alone showed no additional response to smoke extract or BA. Neither smoke extract nor BA affected either cultivar in the dark in the absence of GA_4/7. The partial dormancy-breaking effect of short exposures to red-light was also enhanced by smoke extracts in this highly-dormant cultivar. The results suggest that smoke extracts act in a similar way to cytokinins, by enhancing gibberellin activity in the celery seed system.Abbreviations BA N⁶-benzyladenine - GA_4/7 A₄ and A₇ gibberellin mixture     

Environmental and hormonal regulation of seed dormancy and germination in Cape fynbos Leucospermum R.Br. (Proteaceae) species

The endogenous levels of abscisic acid (ABA), cytokinins (CKs) and gibberellins (GA₁/GA₃ combined) in Leucospermum glabrum embryos were monitored in axes and cotyledons separately during normal germination. Plant growth substance changes were correlated with known morphological, structural and ultrastructural events in the embryo of Proteaceae. The effect of exogenous application of 6-benzyladenine (BA) and GA₄₊₇ under three known dormancy-enforcing environmental conditions were studied in L. glabrum and L. cordifolium. The endogenous levels of the hormone classes GAs and CKs changed phasically during normal germination under a single alternating temperature regime. GA₁/GA₃ levels increased in cotyledons within 3 d of hydration while at the same time initial CK levels decreased. Following this transient peak GAs fell to a low level throughout the germinative period. Subsequently the CKs, Z and ZR, and to a lesser extent their dihydro-derivatives, appeared in both the axes and the cotyledons as fluctuating, transient peaks. Early increases in GAs are thought to control the induction of the germination process. The CK pattern suggests that CKs control at least three major processes of germination sensu stricto following induction: 1) early mobilization of protein and lipid reserves in the axis and later in cotyledons, 2) cotyledon expansion which causes the endotesta to split permitting radicle protrusion and 3) later, radicle growth.Our results indicate that dormancy in intact Leucospermum seeds is enforced by embryo anoxia, regulated by the impermeable exotesta. In addition synthesis of or tissue sensitizing to both hormone classes GAs and CKs depends on moderately low temperature as the primary environmental requirement. For GA synthesis a secondary, daily pulse of high temperature is required. Inhibitory hormones, specifically ABA, appear not to play a role.Abbreviations ABA Abscisic acid - BA 6-benzyladenine - CK Cytokinin - DHZ Dihydrozeatin - DHZR Dihydrozeatin riboside - GA Gibberellin - HPLC High performance liquid chromatography - iP Isopentenyladenine - IPA Isopentenyladenosine - PGS Plant growth substance - RIA Radioimmunoassay - Z Zeatin - ZR Zeatin riboside     

Relationship between gibberellin and cytokinin activity and flowering in Rosa damascena Mill.

Kinetin at 10 mg l^–1 increased the number of flowers produced on Rosa damascena plants while GA₃ inhibited flowering. In the leaves of non-flowering plants GA-like activity was high while specific cytokinin activity (fraction-II) was significantly higher in flowering plants. A novel compound 10- methyldihydrozeatin riboside and isopentenyl-adenine were identified from TLC fraction-II while TLC fraction-I yielded zeatin and 2-hydroxy-6-methylaminopurine.Abbreviations TLC thin layer chromatography - BA N⁶-benzyladenine - GA₃ gibberellic acid CIMAP communication No. 92-40J     

Stimulation of in vitro shoot proliferation from nodal explants of cassava by thidiazuron, benzyladenine and gibberellic acid

Multiple shoots were produced from nodal explants of cassava (Manihot esculenta Crantz) by a two-step procedure: a 6- to 8-day exposure to 0.11–0.22 µM thidiazuron (TDZ) in liquid Murashige and Skoog (MS) medium followed by culture on agar-solidified MS medium supplemented with 2.2 µM 6-benzyladenine (BA) and 1.6 M gibberellic acid (GA₃). TDZ caused the nodal explants to expand and this expansion (growth) continued during culture with BA and GA₃. From this expanded explant, clusters of buds and fasciated stems developed continuously and these gave rise to shoots. The shoot proliferation process was open-ended, yielding an average of 31.5 shoots per nodal explant after 10 weeks of culture with genotype CG 1–56. A positive response was also obtained from seven other genotypes evaluated with this protocol.Abbreviations BA 6-benzyladenine - BM basal medium - DPU 1,3-diphenylurea - GA₃ gibberellie acid - 2iP isopentenyladenine - MSM multiple shoot medium - NAA 1-naphthaleneacetic acid - PGR plant growth regulator - TDZ thidiazuron - Z zeatin     

Influence of plant growth regulators and water stress on ramet induction,rosette engrossment,and fructan accumulation in <Emphasis Type="Italic">Agave tequilana</Emphasis> Weber var. Azul

Agave tequilana Weber var. Azul plants reproduce asexually by producing ramets. Continuous production of ramets throughout the vegetative cycle of the parent delays the time of harvesting of heads for tequila production. Little is known about the factors influencing their emergence. Heads are engrossed rosettes where fructans are stored. We show here that, in plantlets grown in vitro, growth regulators such as 2,4-dichlorophenoxyacetic acid (2,4-D), a combination of 1-naphthaleneacetic acid (NAA)/6-benzyladenine (BA), or abscisic acid (ABA) increased the production of ramets, whereas BA, NAA, gibberellic acid (GA₃), glycerol, or a combination of glycerol/ABA decreased ramet production. Plantlets that developed ramets did not form heads. Head formation was improved on solid media in the presence of BA, NAA, the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC), or the water stress inducer polyethylene glycol (PEG). Basal Murashige–Skoog (MS) liquid media also enhanced rosette engrossment, which was further increased by addition of ACC or PEG. In contrast, CoCl₂, an ethylene biosynthesis inhibitor, reduced rosette engrossment. Furthermore, heads from A. tequilana plantlets grown in tissue culture in MS media, or in MS media supplemented with NAA, ACC or PEG, showed fructan concentrations 10–30 times higher than in leaves from greenhouse-grown plants. Our results indicated that BA, NAA, water stress, and ethylene are critical regulators of rosette engrossment, whereas asexual reproduction in A. tequilana seems to be controlled by a complex hormonal network.     

Plant regeneration from leaf protoplasts of apple

Protoplasts were isolated from young leaves or etiolated shoot apices. For initiation of divisions the protoplasts were embedded in sodium alginate and cultivated in MS or MI medium supplemented with 2.2 M BA, 2.6 M NAA and 2.2 M 2,4-dichlorophenoxyacetic acid. The protoplasts of all seven lines tested developed to protocalluses at high frequencies. No genotypic differences were observed. When BA was used in combination with NAA in the regeneration experiments, only a few protocalluses (highest frequency 3%) exhibited shoot organogenesis. When BA was replaced with thidiazuron, the percentage of protocalluses that developed shoots increased in two of three tested lines to 7% and 56%, respectively. Shoot development was achieved under light conditions. The shoots were then rooted and transferred into soil.Abbreviations ABA abscisic acid - BA 6-benzyladenine - 2,4-d 2,4-dichlorophenoxyacetic acid - FW fresh weight - GA₃ gibberellic acid - IBA indole-3-butyric acid - MES 2-N-morpholinoethane sulphonic acid - NAA -naphthaleneacetic acid     

The effect of plant-hormone pretreatments on ethylene production and synthesis of 1-aminocyclopropane-1-carboxylic acid in water-stressed wheat leaves

Excised wheat (Triticum aestivum L.) leaves, when subjected to drought stress, increased ethylene production as a result of an increased synthesis of 1-aminocyclopropane-1-carboxylic acid (ACC) and an increased activity of the ethyleneforming enzyme (EFE), which catalyzes the conversion of ACC to ethylene. The rise in EFE activity was maximal within 2 h after the stress period, while rehydration to relieve water stress reduced EFE activity within 3 h to levels similar to those in nonstressed tissue. Pretreatment of the leaves with benzyladenine or indole-3-acetic acid prior to water stress caused further increase in ethylene production and in endogenous ACC level. Conversely, pretreatment of wheat leaves with abscisic acid reduced ethylene production to levels produced by nonstressed leaves; this reduction in ethylene production was accompanied by a decrease in ACC content. However, none of these hormone pretreatments significantly affected the EFE level in stressed or nonstressed leaves. These data indicate that the plant hormones participate in regulation of water-stress ethylene production primarily by modulating the level of ACC.Abbreviations ABA abscisic acid - ACC 1-aminocyclopropane-1-carboxylic acid - BA N⁶-benzyladenine - EFE ethylene-forming enzyme - IAA indole-3-acetic acid     

Enhancement of somatic embryogenesis frequency by gibberellic acid in fennel

The effect of GA₃ on somatic embryogenesis from petiole fragments excised from micropropagated fennel plantlets was studied. Explants were maintained for 4 weeks on an induction medium containing, 2,4-d and kinetin and were then transferred to a medium devoid of these growth regulators to allow embryo development. The addition of autoclaved or filter-sterilized GA₃ to the induction medium or to the embryo development medium increased the number of embryogenic explants. No positive effect was observed when GA₃ was added to the micropropagation medium of the mother plantlets. Gibberellic acid also counteracted the inhibiting effect of continuous light on the number of embryogenic explants. Moreover, the embryogenic frequency of petiole explants from several genotypes previously considered as poorly reacting was highly enhanced by GA₃.Abbreviations BA 6-benzyladenine - 2,4-d 2,4-dichlorophenoxyacetic acid - GA₃ Gibberellic acid - IAA Indole-3-acetic acid - IBA Indole-3-butyric acid     

The hormonal control of wheat leaf unrolling

Summary The unrolling of etiolated wheat leaf sections in the dark is stimulated by the application of gibberellic acid (GA₃). GA₃ is most effective if applied for a short time at the beginning of incubation. Kinetin also stimulated leaf unrolling in the dark. AMO1618 and CCC inhibit red light and kinetin-stimulated unrolling. Gibberellin-like substances extracted from red light-treated leaf tissue are effective in stimulating leaf unrolling.Ethylene production in leaf sections is stimulated by IAA, GA₃ and kinetin and inhibited by ABA. A brief exposure to red light decreases the ability of the tissue to produce ethylene. It is concluded that ethylene plays no important role in the control of leaf unrolling by red light or by the application of hormones.Holder of a Science Research Council Studentship.     

Effect of a plant-derived smoke extract, N6-benzyladenine and gibberellic acid on the thermodormancy of lettuce seeds

The effects of a plant-derived smoke extract, BA and GA₃ on the thermodormancy of Grand Rapids lettuce seeds were studied. Thermodormant lettuce seeds treated either with BA, GA₃ or smoke extract alone did not germinate. Combinations of BA with smoke extract and BA with GA₃ were most effective in overcoming induced thermodormancy. GA₃ plus smoke did not break the induced thermodormancy. The effects of the different treatments on germination were concentration dependent. BA was most effective at 10^–5 to 10^–3 M in combination with smoke dilutions 1:5,000 to 1:1,000,000 in overcoming thermodormancy.Abbreviations BA N⁶-benzyladenine; - GA₃ gibberellic acid; - SM smoke extract     

Possible control of gibberellin-induced release of temperature-dependent primary dormancy in seeds of celery (Apium graveolens L.) by transmembrane ion fluxes

The temperature-dependent primary dormancy of cv Florida 683 celery seeds in darkness was broken by GA_4/7 (2 × 10^-4 M) alone but other growth regulators such as BA, ethephon or daminozide were necessary to break dormancy of cv Lathom Blanching seeds in the presence of GA_4/7 at this concentration. Although AgNO₃ partially inhibited both the ethephon- and BA- induced germination of cv Lathom Blanching seeds in the presence of GA_4/7 in the dark it did not affect the promotive action of daminozide. Ethephon did not overcome the inhibitory action of high concentrations of AgNO₃ in the light. The ethylene synthesis inhibitor aminoethoxyvinylglycine (AVG) did not inhibit the germination of cv Lathom Blanching seeds induced by growth regulators in the dark or in the absence of growth regulators in the light. Fusicoccin (FC) did not break celery seed dormancy unless applied in the presence of GA_4/7. Germination of cv Lathom Blanching celery seeds treated with GA_4/7 at 16°C in the dark was inhibited by the K⁺ ionophore benzo-18-crown-C-6 (18-C-6) and in the presence of Ca²⁺ by the Ca²⁺ ionophore A23187; the 18-C-6 inhibition was reversed by BA.It is concluded that the involvement of gibberellin in celery seed dormancy is not dependent on endogenous ethylene and is directly or indirectly controlled through the action of other hormones on transmembrane ion fluxes.     

Somatic embryogenesis and plant regeneration in Quercus acutissima

Immature embryos of Quercus acutissima were collected weekly beginning 5 weeks post-fertilization and cultured on modified MS(Murashige and Skoog) medium containing 1,000 mg/l glutamine and 5 mM proline with different combinations of IBA(0.5–10.0 mg/l) and BA(0 or 1.0 mg/l) in light. The highest percentage of embryogenic cultures occurred on the medium containing 0.5 mg/l IBA or 1.0 mg/l BA and 0.5 mg/l IBA. Four weeks after initiation, the embryogenic cultures were transferred to MS medium without plant growth regulators and cultured for 4 weeks. The somatic embryos were then transferred to germination medium. The best germination results were achieved from WPM(Woody Plant Medium) containing 0.1 mg/l BA. Plantlets from somatic embryos were incubated on WPM supplemented with 0.2 mg/l BA for 4 weeks and plantlets with well developed shoots and roots were transplanted to perlite and peat moss(11, v/v) mixtures and placed in a culture room. After being hardened off for 8 weeks, they were transferred outdoors where they grew.Abbreviation BA N⁶-benzyladenine - IBA indole-3-butyric acid - GA₃ gibberellic acid - ABA abscisic acid - MS Murashige & Skoog Medium - WPM Woody Plant medium     

Interaction of jasmonic acid with some plant growth regulators in the control of apple (Malus domestica) embryo germination

Embryos isolated from dormant apple seeds were treated with jasmonic acid (JA), gibberellin A₃ (GA₃), abscisic acid (ABA) and hydrogen cyanide in darkness and in light. The chemicals were present in the culture medium continuously and simultaneously or applied for 2 days and in different sequences. All treatments stimulated embryo germination except ABA, which was strongly inhibitory. Additive effects of JA with light and with GA₃ on embryo germination were observed, whereas ABA interacted synergically with JA, HCN and light. ABA and GA₃ were most effective when applied early during embryo incubation, but the late JA treatment was more stimulatory. It is concluded that JA does not act on the regulatory pathway that is initiated by light and which leads to embryo germination through gibberellin accumulation and alkaline lipase activation. ABA and HCN appear to be involved in the control of this pathway. JA and ABA may be involved in the control of alkaline lipase activity, independently of this regulatory chain.Abbreviations ABA abscisic acid - GA₃ gibberellin A₃ - JA jasmonic acid     

Relationships between position on the parent plant and germination characteristics of seeds of parsley (Petroselinium crispum Nym.)

The percentage germination of seeds of parsley cv. Imperial Curled was higher in the light than in the dark, the high temperature limits for germination being 30 and 28°C for light and dark respectively. At the higher temperatures, the germination rate was slower in the dark. At 30°C, treatment with a gibberellin A_4/7 mixture at 2 × 10^–4 M partially alleviated the inhibiting effect of darkness on the germination percentage. Pre-incubation of parsley seeds at 35°C in the dark for 30 h increased the rate, but decreased the percentage, of germination of seeds incubated at 15°C in the light. Germination and seedling emergence studies were made on seed harvested from four different umbel positions. Although heavier seeds were produced from primary umbels than from other umbel orders, they were less viable as measured by seedling emergence in the glasshouse. The rate of emergence was decreased with increasing umbel order i.e. with later seed development: this was reflected in subsequent seedling weights, with seedlings from quarternary umbel seeds being about half the weight of those from primary umbel seeds. The upper temperature limit for dark germination was only slightly affected by umbel order, with quarternary umbel seeds being the most thermo-inhibited.Abbreviations BA N⁶-benzyladenine - GA_4/7 a mixture of gibberellins A₄ and A₇ - SD8339 6-benzyl-amino-9-(tetrahydropyran-2-yl)-9H-purine     

Integrative regulation of leaf morphogenesis by gibberellic and abscisic acids in the aquatic angiosperm proserpinaca palustris L

A two-hormone system regulating leaf development in the heterophyllous amphibious angiosperm Proserpinaca palustris L. is described. Aerial shoots develop expanded, lanceolate, serrate leaves under long-day photoperiods (LD, 16 h light: 8 h dark), whereas growth under short days (SD, 10 h light: 14 h dark) induces dissected leaf formation. The photoperiodic effect on leaf development of aerial shoots involves changes in endogenous gibberellins (GAs) since plants grown under SD in the presence of GA₃ develop expanded lanceolate serrate leaves. However, when submerged, shoots develop highly dissectedaquatic leaves regardless of photoperiod or GA₃ treatment. In the present study, submerged plants exposed to 1.0 or 5.0 μM abscisic acid (ABA) developed aerial-type leaves typical of the photoperiod under which they were cultured. Both exogenous ABA (5.0 μM) and GA₃ (10 μM) treatments were required for laminar expansion to occur on submerged shoots under SD. It is suggested that (1) leaf development in Proserpinaca is regulated by both endogenous GAs and ABA, and (2) the endogenous status of these phytohormones is modulated by different environmental stimuli of photoperiod and water stress, respectively. The adaptive significance of this mechanism is discussed.