Analyses of abscisic acid (ABA), ent-kaurenoids and gibberellins (GAs) showed that there were major changes in the contents of these compounds associated with germination of after-ripened barley (Hordeum vulgare cv. Schooner and cv. Proctor) grain but not in hydrated dormant grain. Embryos from dormant and after-ripened dry grain contained similar amounts of ABA, of ent-kaurenoids and of GAs, determined by gas chromatography-mass spectrometry-selected ion monitoring. In embryos of after-ripened grain, ABA content decreased rapidly after hydration and ABA appeared to be metabolized (inactivated) to phaseic acid (PA) rather than diffusing into the endosperm or the surrounding medium as previously thought. Similar changes in ABA occurred in hydrated dormant grain during germination in darkness. Accumulation of ent-kaurenoids and GAs, including GA1, the first biologically active GA in the early 13-hydroxylation biosynthetic pathway, occurred to a much greater extent in after-ripened than in dormant grain and these changes occurred mainly after 18 h of hydration when ABA had already decreased and germination was occurring. The block in ent-kaurenoid and GA synthesis in dormant grain appeared to occur prior to ent-kaurene in the biosynthetic pathway. These results are consistent with the view that ABA is the primary effector of dormancy and that after-ripening involves the development of the ability to reduce the amount of ABA quickly following hydration. Accumulation of GAs does not appear to be causally related to loss of dormancy but it does appear to be related to germination.
An attempt has been made to uncouple the effects of the two primary components of shade light, a reduced red to far-red (R/FR) ratio and low photosynthetically active radiation (PAR), on the elongation of the youngest internode of sunflower (Helianthus annuus) seedlings. Maximal internode growth (length and biomass) was induced by a shade light having a reduced R/FR ratio (0.85) under the low PAR of 157 micromol m(-2) s(-1). Reducing the R/FR ratio under normal PAR (421 micromol m(-2) s(-1)) gave similar growth trends, albeit with a reduced magnitude of the response. Leaf area growth showed a rather different pattern, with maximal growth occurring at the higher (normal) PAR of 421 micromol m(-2) s(-1)), but with variable effects being seen with changes in light quality. Reducing the R/FR ratio (by enrichment with FR) gave significant increases in gibberellin A(1) (GA(1)) and indole-3-acetic acid (IAA) contents in both internodes and leaves. By contrast, a lower PAR irradiance had no significant effect on GA(1) and IAA levels in internodes or leaves, but did increase the levels of other GAs, including two precursors of GA(1). Interestingly, both leaf and internode hormone content (GAs, IAA) are positively and significantly correlated with growth of the internode, as are leaf levels of abscisic acid (ABA). However, changes in these three hormones bear little relationship to leaf growth. By implication, then, the leaf may be the major source of GAs and IAA, at least, for the rapidly elongating internode. Several other hormones were also assessed in leaves for plants grown under varying R/FR ratios and PARs. Leaf ethylene production was not influenced by changes in R/FR ratio, but was significantly reduced under the normal (higher) PAR, the irradiance treatment which increased leaf growth. Levels of the growth-active free base and riboside cytokinins were significantly increased in leaves under a reduced R/FR ratio, but only at the higher (normal) PAR irradiance; other light quality treatments evoked no significant changes. Taken in toto, these results indicate that both components of shade light can influence the levels of a wide range of endogenous hormones in internodes and leaves while evoking increased internode elongation and biomass accumulation. However, it is light quality changes (FR enrichment) which are most closely tied to increased hormone content, and especially with increased GA and IAA levels. Finally, the increases seen in internode and leaf GA content with a reduced R/FR ratio are consistent with FR enrichment inducing an overall increase in sunflower seedling GA biosynthesis.
The gibberellin-deficient mutant, ga1-1 (NG5) of Arabidopsis thaliana, when induced by 16-h-long days, will form floral buds. However, the flower stalk is very short and floral organs within the flowers remain undeveloped; petal growth is arrested, with the petals being scaly and translucent, the stamens are abortive, the filaments do not elongate, and the pollen does not mature. Sepals and pistils are also underdeveloped. All of the above defects of this mutant can be completely eliminated if certain gibberellins (GAs) are applied to the young floral buds. That is, the applied GA acts to normalize not only plant height but also development of floral organs, thereby yielding good seed set from self-pollination. There were appreciable differences in the efficacy of different GA structures in normalizing the various floral organs. For seed production, the order of efficacy was 2,2-dimethyl GA4 > GA7 > GA3 = GA4 > GA1 > GA5 = GA9. When 2,2-dimethyl GA4 was used to determine an optimal GA dose, the following pattern emerged: filament elongation and pollen development, 1-10 ng; petal and pistil growth, 1 ng; sepal growth, 0.1 ng; papilla elongation, 0.01 ng. However, one application at these doses was insufficient to normalize the flowers, which were formed one after another, and a continuing supply of GA at the optimal dose was required for normal flower development and seed set. We conclude from this work that GAs play an essential role in the development of floral organs of Arabidopsis and that petals and stamens (filaments and pollen) in particular develop normally only when GAs are present at the optimal level.Key words: Arabidopsis thaliana, floral organ development, gibberellin, gibberellin-deficient mutant, petal and pollen development, reproductive function.
Plants subjected to abiotic stresses such as extreme high and low temperatures, drought or salinity, often exhibit decreased vegetative growth and reduced reproductive capabilities. This is often associated with decreased photosynthesis via an increase in photoinhibition, and accompanied by rapid changes in endogenous levels of stress-related hormones such as abscisic acid (ABA), salicylic acid (SA) and ethylene. However, certain plant species and/or genotypes exhibit greater tolerance to abiotic stress because they are capable of accumulating endogenous levels of the zwitterionic osmolyte-glycinebetaine (GB). The accumulation of GB via natural production, exogenous application or genetic engineering, enhances plant osmoregulation and thus increases abiotic stress tolerance. The final steps of GB biosynthesis occur in chloroplasts where GB has been shown to play a key role in increasing the protection of soluble stromal and lumenal enzymes, lipids and proteins, of the photosynthetic apparatus. In addition, we suggest that the stress-induced GB biosynthesis pathway may well serve as an additional or alternative biochemical sink, one which consumes excess photosynthesis-generated electrons, thus protecting photosynthetic apparatus from overreduction. Glycinebetaine biosynthesis in chloroplasts is up-regulated by increases in endogenous ABA or SA levels. In this review, we propose and discuss a model describing the close interaction and synergistic physiological effects of GB and ABA in the process of cold acclimation of higher plants.
Gibberellins A1, A3, and iso-A3 were identified from aseptic cultures of Azospirillum lipoferum strain op 33 by capillary gas chromatography-mass spectrometry (GC-MS) and GC-MS-selected ion monitoring. There were 20 to 40 picograms (in GA3 equivalents, estimated from bioassay) of gibberellins A1 and A3 per milliliter of cell culture (containing 10' cells).
Structural requirements for florigenic activity among gibberellins (GAs) and GA derivatives, including several new ones, applied once to leaves of Lolium temulentum, were examined. The compounds were applied to plants kept either in non-inductive short days (SD) or exposed to one inductive long day (LD). Inflorescence initiation and stem-elongation responses were assessed three weeks later. Among the GAs used, the range in effective dose for inflorescence initiation was more than 1000-fold, but substantially less for stem elongation. Some GAs promoted both stem elongation and inflorescence initiation, some promoted one without the other, and some affected neither. The structural features enhancing florigenic activity were often different from those enhancing stem elongation. Except in the case of 2,2-dimethyl GA4, a double bond in the A ring at either C-1,2 or C-2,3 was essential for high florigenic activity, though not for stem elongation. A free carboxy group was needed for both. Inflorescence initiation in Lolium was enhanced by hydroxylation at C-12, -13 and -15, whereas hydroxylation at C-3 reduced the effect on inflorescence initiation but increased that on stem elongation. A 12β-hydroxyl was more effective than the α epimer for inflorescence initiation whereas the reverse was true for stem elongation. Although such differential effectiveness of GAs for inflorescence initiation and for stem elongation could reflect differences in uptake, transport or metabolism, we suggest that it is indicative of specific structural requirements for inflorescence initiation.
Snow tussocks (Chionochloa spp.) in New Zealand exhibit extreme mast seeding which is synchronised within and among populations and species over large spatial scales. This masting behaviour satiates three endemic insect seed predators, and the weather cue that triggers heavy flowering has been reported as a very warm austral summer in the year before flowering. Here we elucidate the details of flowering, predator satiation, and temperature cues for Chionochloa. A 22-year observational data set for Chionochloa pallens from 1,070 m elevation on Mt Hutt, Canterbury, indicates that flowering was highly variable (CV = 1.79) which was effective in predator satiation, and the key cue for heavy flowering was warm temperatures between 1 January and 7 February the year before flowering. Both the number of inflorescences per tussock (the most variable component of reproductive output) and the number of florets per inflorescence were increased. Surprisingly, there were also same-season effects of temperatures between October and December on the number of inflorescences per tussock produced in late December through February. Experimental warming of tussocks using open-topped clear plastic enclosure tubes verified the sameseason observations by significantly increasing flowering in the current season, but much less often increased flowering in the subsequent season. A comparison of masting patterns 1990-2008 with altitude in two tussock species, C. pallens and C. macra, indicates that at higher altitudes (ca. 1,580 m vs. 1,070 m) the temperature thresholds for a given flowering effort were lowered. The lowering of thresholds (2.1°C in C. pallens and 1.5°C in C. macra) was similar to the reduction in mean growing-season air temperatures (2.0°C) found between the sites. Thus, despite different local temperatures at high and low altitude sites, the different populations flowered with a similar intensity each year. The primary benefit of mast seeding in Chionochloa appears to be predator satiation, and the key predator (the cecidomyiid Eucalyptodiplosis chionochloae) has extended diapause. If diapause can be influenced by climate cues, the exact nature of the climate cues will be important to the evolutionary interaction between the plant and its seed predators.
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