Abscisic acid (ABA) was discovered in the 1950s to be a phytohormone affecting leaf abscision and bud dormancy. It was soon characterized as a sesquiterpene derived from mevalonate although certain steps of its biosynthesis in plants are still unknown (Li and Walton, 1987; Zeevaart and Creelman, 1988). Continuing work on ABA has shown that it mediates various developmental and physiological processes that affect the agronomic performance of crop plants (Austin et al., 1982; Ramagopal, 1987). These proc? esses include embryo maturation and germination as well as the response of vegetative tissues to osmotic stress (Singh et al., 1987; Zeevaart and Creelman, 1988). ABA levels increase in tissues subjected to osmotic stress by desiccation, salt, or cold (Henson, 1984; Mohapatra et al., 1988). Under these conditions, specific genes are ex? pressed that can also be induced in unstressed tissues by the application of exogenous ABA (Singh et al., 1987; Gomez et al., 1988; Mundy and Chua, 1988). Some of these genes are also expressed during the normal embryogenic program when seeds desiccate and embryos be? come dormant (Dure et al., 1981). Although different sets of ABA-responsive genes exhibit different patterns of de? velopmental and tissue-specific expression, some of them appear to be part of a general reaction to osmotic stress. This system is a normal part of the embryogenic program but is inducible in vegetative tissues at other times in the plant life cycle. Several ABA-responsive genes have now been isolated (Baker et al., 1988; Gomez et al., 1988; Marcotte et al., 1988; Mundy and Chua, 1988; Vilardell et al., 1990; Yamaguchi-Shinozaki et al., 1990). A major goal of the research discussed below is to understand the role these genes play in osmotic stress and desiccation tolerance.

Gene expression in response to abscisic acid and osmotic stress.

Lodging-resistant rice (Oryza sativa) cultivars usually show slow grain filling when nitrogen is applied in large amounts. This study investigated the possibility that a hormonal change may mediate the effect of water deficit that enhances whole plant senescence and speeds up grain filling. Two rice cultivars showing high lodging resistance and slow grain filling were field grown and applied with either normal or high amount nitrogen (HN) at heading. Well-watered and water-stressed (WS) treatments were imposed 9 days post anthesis to maturity. Results showed that WS increased partitioning of fixed (14)CO(2) into grains, accelerated the grain filling rate but shortened the grain filling period, whereas the HN did the opposite way. Cytokinin (zeatin + zeatin riboside) and indole-3-acetic acid contents in the grains transiently increased at early filling stage and WS treatments hastened their declines at the late grain filling stage. Gibberellins (GAs; GA(1) + GA(4)) in the grains were also high at early grain filling but HN enhanced, whereas WS substantially reduced, its accumulation. Opposite to GAs, abscisic acid (ABA) in the grains was low at early grain filling but WS remarkably enhanced its accumulation. The peak values of ABA were significantly correlated with the maximum grain filling rates (r = 0.92**, P < 0.01) and the partitioning of fixed (14)C into grains (r = 0.95**, P < 0.01). Exogenously applied ABA on pot-grown HN rice showed similar results as those by WS. Results suggest that an altered hormonal balance in rice grains by water stress during grain filling, especially a decrease in GAs and an increase in ABA, enhances the remobilization of prestored carbon to the grains and accelerates the grain filling rate.

Hormonal Changes in the Grains of Rice Subjected to Water Stress during Grain Filling

The GAs (gibberellins) comprise a large group of diterpenoid carboxylic acids that are ubiquitous in higher plants, in which certain members function as endogenous growth regulators, promoting organ expansion and developmental changes. These compounds are also produced by some species of lower plants, fungi and bacteria, although, in contrast to higher plants, the function of GAs in these organisms has only recently been investigated and is still unclear. In higher plants, GAs are synthesized by the action of terpene cyclases, cytochrome P450 mono-oxygenases and 2-oxoglutarate-dependent dioxygenases localized, respectively, in plastids, the endomembrane system and the cytosol. The concentration of biologically active GAs at their sites of action is tightly regulated and is moderated by numerous developmental and environmental cues. Recent research has focused on regulatory mechanisms, acting primarily on expression of the genes that encode the dioxygenases involved in biosynthesis and deactivation. The present review discusses the current state of knowledge on GA metabolism with particular emphasis on regulation, including the complex mechanisms for the maintenance of GA homoeostasis.

Gibberellin biosynthesis and its regulation.

▪ Abstract The recent impressive progress in research on gibberellin (GA) biosynthesis has resulted primarily from cloning of genes encoding biosynthetic enzymes and studies with GA-deficient and GA-insensitive mutants. Highlights include the cloning of ent-copalyl diphosphate synthase and ent-kaurene synthase (formally ent-kaurene synthases A and B) and the demonstration that the former is targeted to the plastid; the finding that the Dwarf-3 gene of maize encodes a cytochrome P450, although of unknown function; and the cloning of GA 20-oxidase and 3β-hydroxylase genes. The availability of cDNA and genomic clones for these enzymes is enabling the mechanisms by which GA concentrations are regulated by environmental and endogenous factors to be studied at the molecular level. For example, it has been shown that transcript levels for GA 20-oxidase and 3β-hydroxylase are subject to feedback regulation by GA action and, in the case of the GA 20-oxidase, are regulated by light. Also discussed is other new info...

GIBBERELLIN BIOSYNTHESIS: Enzymes, Genes and Their Regulation.

The phytohormone abscisic acid is probably present in all higher plants. This hormone is necessary for regulation of several events during seed development and for the response to environmental stresses such as desiccation, salt and cold. An important part of the physiological response to abscisic acid is achieved through gene expression. Here, we summarize the current knowledge of regulation of abscisic acid-induced transcription. The main focus is on a description of the known abscisic acid-responsive cis-elements, their properties and the possible transacting factors binding to the elements. Results have shown that cooperative action of cis-elements and the promoter configuraton is crucial for regulation by abscisic acid. Furthermore, several elements are organ- and species-specific. Recent studies of the chromatin structure of abscisic acid-responsive genes point to the importance of induction of transcription by coactivators or by phosphorylation/dephosphorylation of transcription factors. An interesting example of activation by a cofactor is the cooperative action between abscisic acid-signaling and the regulatory protein Viviparous 1 through the abscisic acid responsive element.

Regulation of abscisic acid-induced transcription.

Changes in the kind and level of endogenous gibberellins (GAs) in the shoot and ear of rice (Oryza sativa L. japonica variety, cultivar Nihonbare) through its life cycle were investigated. Gibberellin A19 (GA19) was the major GA, and GA1, GA20, GA29 and GA53 were minor GAs in the shoot of rice. Besides these 13-hydroxylated (13-OH) GAs, two non-13-hydroxylated (13-H) GAs, namely GA4 and GA34, were identified in the shoot (containing the ear) collected at the heading stage.Further studies on the distribution and changes in the level of GAs in reproductive organs revealed that GA4 was highly concentrated in anthers collected just before anthesis, but was not detected in immature seeds collected about a week after anthesis. These results suggest that 13-H GAs may be biosynthesized in specific organs at a specific stage of the life cycle, and that they may be concerned with the regulation of reproductive growth, while vegetative growth is regulated by 13-OH GAs, probably GA1.

https://www.jstage.jst.go.jp/article/bbb1961/52/5/52_5_1189/_pdf

Fluctuation and Localization of Endogenous Gibberellins in Rice

The major endogenous gibberellin (GA) in shoots, roots and ears of the rice plant, Oryza sativa L. japonica cv. Nihonbare, was identified as GA19 by combined gas liquid chromatography-mass spectrometry (GC-MS) and GC-selected ion current monitoring (GC-SICM). Another GA present in these tissues in small quantity was tentatively identified as GA1 by GC-SICM, and GA4 may be present in the seeds (kernels) of 3rd-leaf-stage seedlings. Using GC-SICM, the GA19 content was quantified throughout the life cycle of rice plants. It was found to reach high levels (ca. 10–15 μg/kg fresh weight) in 3rd-leaf seedlings, at panicle initiation (shoots), and during heading and anthesis (ears). The levels of GA19 in Oryza sativa indica cv. T-136 underwent changes closely similar to those found in Nihonbare. The growth-promoting activity in rice of exogenous GA19 is generally considerably less than that of GA1. It therefore seems possible that GA19 functions as a “pool GA”. The level of active GAs such as GA1 may be regulated by the rate of biosynthesis of GA19 or its metabolic conversions.

Identification of gibberellins in the rice plant and quantitative changes of gibberellin A19 throughout its life cycle.

Seasonal Changes of GA1, GA19 and Abscisic Acid in Three Rice Cultivars

The change in the levels of endogenous cytokinins in the shoot and grain of rice (Oryza sativa L. japonica cultivar Nihonbare) during its growth and development was followed by selected ion monitoring using deuterium-labelled internal standards. Five cytokinins, zeatin (Z), cis-Z, ribosyl-zeatin (RZ), cis-RZ and 6-(3-methyl-but-2-enylamino)purine riboside were detected in the shoot, root and ear of rice. The maximum contents of RZ and Z in the shoot and root were found at the maximum tillering stage. In the ear, the levels of Z and RZ changed markedly. It is therefore suspected that RZ and Z play some roles in the tillering and seed growth in rice.

Fluctuation of endogenous cytokinin contents in rice during its life cycle ― quantification of cytokinins by selected ion monitoring using deuterium-labelled internal standards

Yasuo Ota

Papers

Fluctuation and Localization of Endogenous Gibberellins in Rice

Identification of gibberellins in the rice plant and quantitative changes of gibberellin A19 throughout its life cycle.

Seasonal Changes of GA1, GA19 and Abscisic Acid in Three Rice Cultivars

Fluctuation of endogenous cytokinin contents in rice during its life cycle ― quantification of cytokinins by selected ion monitoring using deuterium-labelled internal standards

Identification of Cytokinins in Root Exudate of the Rice Plant