The level of abscisic acid (ABAabscisic acid) in any particular tissue in a plant is determined by the rate of biosynthesis and catabolism of the hormone. Therefore, identifying all the genes involved in the metabolism is essential for a complete understanding of how this hormone directs plant growth and development. To date, almost all the biosynthetic genes have been identified through the isolation of auxotrophic mutants. On the other hand, among several ABA catabolic pathways, current genomic approaches revealed that Arabidopsis CYP707A genes encode ABA 8′-hydroxylases, which catalyze the first committed step in the predominant ABA catabolic pathway. Identification of ABA metabolic genes has revealed that multiple metabolic steps are differentially regulated to fine-tune the ABA level at both transcriptional and post-transcriptional levels. Furthermore, recent ongoing studies have given new insights into the regulation and site of ABA metabolism in relation to its physiological roles.

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Abscisic acid biosynthesis and catabolism

This review focuses mainly on eudicot seeds, and on the interactions between abscisic acid (ABA), gibberellins (GA), ethylene, brassinosteroids (BR), auxin and cytokinins in regulating the interconnected molecular processes that control dormancy release and germination. Signal transduction pathways, mediated by environmental and hormonal signals, regulate gene expression in seeds. Seed dormancy release and germination of species with coat dormancy is determined by the balance of forces between the growth potential of the embryo and the constraint exerted by the covering layers, e.g. testa and endosperm. Recent progress in the field of seed biology has been greatly aided by molecular approaches utilizing mutant and transgenic seeds of Arabidopsis thaliana and the Solanaceae model systems, tomato and tobacco, which are altered in hormone biology. ABA is a positive regulator of dormancy induction and most likely also maintenance, while it is a negative regulator of germination. GA releases dormancy, promotes germination and counteracts ABA effects. Ethylene and BR promote seed germination and also counteract ABA effects. We present an integrated view of the molecular genetics, physiology and biochemistry used to unravel how hormones control seed dormancy release and germination.

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Plant hormone interactions during seed dormancy release and germination

Plants are exposed to a variety of abiotic stresses in nature and exhibit unique and complex responses to these stresses depending on their degree of plasticity involving many morphological, cellular, anatomical, and physiological changes. Phytohormones are known to play vital roles in the ability of plants to acclimatize to varying environments, by mediating growth, development, source/sink transitions and nutrient allocation. These signal molecules are produced within the plant, and also referred as plant growth regulators. Although plant response to salinity depends on several factors; nevertheless, phytohormones are thought to be the most important endogenous substances that are critical in modulating physiological responses that eventually lead to adaptation to salinity. Response usually involves fluctuations in the levels of several phytohormones, which relates with changes in expression of genes involved in their biosynthesis and the responses they regulate. Present review described the potential role of different phytohormones and their balances against salinity stress and summarized the research progress regarding plant responses towards salinity at physiological and molecular levels. We emphasized the role of abscisic acid, indole acetic acid, cytokinins, gibberellic acid, salicylic acid, brassinosteroids, jasmonates, ethylene and triazoles in mediating plant responses and discussed their crosstalk at various baseline pathways transduced by these phytohormones under salinity. Current progress is exemplified by the identification and validation of several significant genes that enhanced crops tolerance to salinity, while missing links on different aspects of phytohormone related salinity tolerance are pointed out. Deciphering mechanisms by which plant perceives salinity and trigger the signal transduction cascades via phytohormones is vital to devise salinity related breeding and transgenic approaches.

Phytohormones and plant responses to salinity stress: a review

Endogenous abscisic acid (ABA) levels are regulated by both biosynthesis and catabolism of the hormone. ABA 8′-hydroxylase is considered to be the key catabolic enzyme in many physiological processes. We have previously identified that four members of the Arabidopsis (Arabidopsis thaliana) CYP707A gene family (CYP707A1 to CYP707A4) encode ABA 8′-hydroxylases, and that the cyp707a2 mutants showed an increase in ABA levels in dry and imbibed seeds. In this study, we showed that the cyp707a1 mutant accumulated ABA to higher levels in dry seeds than the cyp707a2 mutant. Expression analysis showed that the CYP707A1 was expressed predominantly during mid-maturation and was down-regulated during late-maturation. Concomitantly, the CYP707A2 transcript levels increased from late-maturation to mature dry seed. Phenotypic analysis of single and double cyp707a mutants indicates that the CYP707A1 is important for reducing ABA levels during mid-maturation. On the other hand, CYP707A2 is responsible for the regulation of ABA levels from late-maturation to germination. Moreover, CYP707A1 and CYP707A3 were also shown to be involved in postgermination growth. Spatial expression analysis suggests that CYP707A1 was expressed predominantly in embryo during mid-maturation, whereas CYP707A2 expression was detected in both embryo and endosperm from late-maturation to germination. Our results demonstrate that each CYP707A gene plays a distinct role during seed development and postgermination growth.

CYP707A1 and CYP707A2, Which Encode Abscisic Acid 8′-Hydroxylases, Are Indispensable for Proper Control of Seed Dormancy and Germination in Arabidopsis

Plants are sessile beings, so the need of mechanisms to flee from unfavorable circumstances has provided the development of unique and sophisticated responses to environmental stresses. Depending on the degree of plasticity, many morphological, cellular, anatomical, and physiological changes occur in plants in response to abiotic stress. Phytohormones are small molecules that play critical roles in regulating plant growth and development, as well as stress tolerance to promote survival and acclimatize to varying environments. To congregate the challenges of salinity, temperature extremes, and osmotic stress, plants use their genetic mechanism and different adaptive and biological approaches for survival and high production. In the present attempt, we review the potential role of different phytohormones and plant growth-promoting rhizobacteria in abiotic stresses and summarize the research progress in plant responses to abiotic stresses at physiological and molecular levels. We emphasized the regulatory circuits of abscisic acid, indole acetic acid, cytokinins, gibberellic acid, salicylic acid, brassinosteroids, jasmonates, ethylene, and triazole on exposure to abiotic stresses. Current progress is exemplified by the identification and validation of several significant genes that enhanced crop tolerance to stress in the field. These findings will make the modification of hormone biosynthetic pathways for the transgenic plant generation with augmented abiotic stress tolerance and boosting crop productivity in the coming decades possible.

Potential role of phytohormones and plant growth-promoting rhizobacteria in abiotic stresses: consequences for changing environment.

Germination of lettuce (Lactuca sativa L. cv. 'Grand Rapids') seeds was inhibited at high temperatures (thermoinhibition). Thermoinhibition at 28 degrees C was prevented by the application of fluridone, an inhibitor of abscisic acid (ABA) biosynthesis. At 33 degrees C, the sensitivity of the seeds to ABA increased, and fluridone on its own was no longer effective. However, a combined application of fluridone and gibberellic acid (GA3) was able to restore the germination. Exogenous GA3 lowered endogenous ABA content in the seeds, enhancing catabolism of ABA and export of the catabolites from the intact seeds. The fluridone application also decreased the ABA content. Consequently, the combined application of fluridone and GA3 decreased the ABA content to a sufficiently low level to allow germination at 33 degrees C. There was no significant temperature-dependent change in endogenous GA1 contents. It is concluded that ABA is an important factor in the regulation of thermoinhibition of lettuce seed germination, and that GA affects the temperature responsiveness of the seeds through ABA metabolism.

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Shusuke Kawahara

Papers

Abscisic acid in the thermoinhibition of lettuce seed germination and enhancement of its catabolism by gibberellin