Abscisic acid (ABA) is a plant hormone that regulates seed dormancy and germination. Seeds undergo changes in both ABA content and sensitivity during seed development and germination in response to internal and external cues. Recent advances in functional genomics have revealed the integral components involved in ABA metabolism (biosynthesis and catabolism) and perception, the core signalling pathway, as well as the factors that trigger ABA-mediated transcription. These allow for comparative studies to be conducted on seeds under different environmental conditions and from different genetic backgrounds. This review summarizes our understanding of the control of ABA content and the responsiveness of seeds to afterripening, light, high temperature and nitrate, with a focus on which tissues are involved in its metabolism and signalling. Also described are the regulators of ABA metabolism and signalling, which potentially act as the node for hormone crosstalk. Integration of such knowledge into the complex and diverse events occurring during seed germination will be the next challenge, which will allow for a clearer understanding of the role of ABA.

The regulation of abscisic acid (ABA) biosynthesis is essential for plant responses to drought stress. In this study, we examined the tissue-specific localization of ABA biosynthetic enzymes in turgid and dehydrated Arabidopsis (Arabidopsis thaliana) plants using specific antibodies against 9-cis-epoxycarotenoid dioxygenase 3 (AtNCED3), AtABA2, and Arabidopsis aldehyde oxidase 3 (AAO3). Immunohistochemical analysis revealed that in turgid plants, AtABA2 and AAO3 proteins were localized in vascular parenchyma cells most abundantly at the boundary between xylem and phloem bundles, but the AtNCED3 protein was undetectable in these tissues. In water-stressed plants, AtNCED3 was detected exclusively in the vascular parenchyma cells together with AtABA2 and AAO3. In situ hybridization using the antisense probe for AtNCED3 showed that the drought-induced expression of AtNCED3 was also restricted to the vascular tissues. Expression analysis of laser-microdissected cells revealed that, among nine drought-inducible genes examined, the early induction of most genes was spatially restricted to vascular cells at 1 h and then some spread to mesophyll cells at 3 h. The spatial constraint of AtNCED3 expression in vascular tissues provides a novel insight into plant systemic response to drought stresses.

In angiosperms, a double fertilization event initiates the development of two distinct structures, the embryo and endosperm. The endosperm plays an important role in supporting embryonic growth by supplying nutrients, protecting the embryo and controlling embryo growth by acting as a mechanical barrier during seed development and germination. Its structure and function in the mature dry seed is divergent and specialized among different plant species. A subset of endospermic tissues are composed of living cells even after seed maturation, and play an active role in the regulation of seed germination. Transcriptome analysis has provided new insights into the regulatory functions of the endosperm during seed germination. It is well known that the embryo secretes signals to the endosperm to induce the degradation of the seed reserve and to promote endosperm weakening during germination. Recent advances in seed biology have shown that the endosperm is capable of sensing environmental signals, and can produce and secrete signals to regulate the growth of the embryo. Thus, germination is a systemic response that involves bidirectional interactions between the embryo and endosperm.

Levels of endogenous abscisic acid (ABA) are changed dynamically in response to environmental conditions. The ABA 8'-hydroxylase is a key enzyme in ABA catabolism and is encoded by CYP707A genes. In this study, we examined physiological roles of Arabidopsis (Arabidopsis thaliana) CYP707As in the plant's response to changes in humidity. The cyp707a1 and cyp707a3 mutants displayed lower stomatal conductance under turgid conditions (relative humidity 60%) than the wild type. When wild-type plants were transferred to high-humidity conditions (relative humidity 90%), CYP707A1 and CYP707A3 transcript levels increased, followed by the reduction of ABA levels. The cyp707a3 mutant exhibited high ABA levels even after transferring to high-humidity conditions, whereas, under similar conditions, the cyp707a1 mutant exhibited low ABA levels comparable to the wild type. Analysis of spatial expression patterns by using transgenic plants harboring a promoterbeta-glucuronidase gene indicated that high-humidity-induced expression of CYP707A1 and CYP707A3 occurred primarily in guard cells and vascular tissues, respectively. Furthermore, stomatal closure of the cyp707a1 mutant, but not cyp707a3 mutant, was ABA hypersensitive when epidermal peel was treated with exogenous ABA, suggesting that CYP707A1 is essential for ABA catabolism inside the guard cells. These results implicate that CYP707A3 reduces the amount of mobile ABA in vascular tissues in response to high humidity, whereas CYP707A1 inactivates local ABA pools inside the guard cells. Taken together, ABA catabolism in both vascular tissues and guard cells participates in the systemic ABA action that controls stomatal movement in response to high humidity.

Abscisic acid (ABA) regulates various aspects of plant physiology, including promoting seed dormancy and adaptive responses to abiotic and biotic stresses. In addition, ABA plays an im-portant role in growth and development under non-stressed conditions. This review summarizes phenotypes of ABA biosynthesis and signaling mutants to clarify the roles of basal ABA in growth and development. The promotive and inhibitive actions of ABA in growth are characterized by stunted and enhanced growth of ABA-deficient and insensitive mutants, respectively. Growth regulation by ABA is both promotive and inhibitive, depending on the context, such as concentrations, tissues, and environmental conditions. Basal ABA regulates local growth including hyponastic growth, skotomorphogenesis and lateral root growth. At the cellular level, basal ABA is essential for proper chloroplast biogenesis, central metabolism, and expression of cell-cycle genes. Basal ABA also regulates epidermis development in the shoot, by inhibiting stomatal development, and deposition of hydrophobic polymers like a cuticular wax layer covering the leaf surface. In the root, basal ABA is involved in xylem differentiation and suberization of the endodermis. Hormone crosstalk plays key roles in growth and developmental processes regulated by ABA. Phenotypes of ABA-deficient and insensitive mutants indicate prominent functions of basal ABA in plant growth and development.

Seeds respond to multiple different environmental stimuli that regulate germination. Nitrate stimulates germination in many plants but how it does so remains unclear. Here we show that the Arabidopsis NIN-like protein 8 (NLP8) is essential for nitrate-promoted seed germination. Seed germination in nlp8 loss-of-function mutants does not respond to nitrate. NLP8 functions even in a nitrate reductase-deficient mutant background, and the requirement for NLP8 is conserved among Arabidopsis accessions. NLP8 reduces abscisic acid levels in a nitrate-dependent manner and directly binds to the promoter of CYP707A2, encoding an abscisic acid catabolic enzyme. Genetic analysis shows that NLP8-mediated promotion of seed germination by nitrate requires CYP707A2. Finally, we show that NLP8 localizes to nuclei and unlike NLP7, does not appear to be activated by nitrate-dependent nuclear retention of NLP7, suggesting that seeds have a unique mechanism for nitrate signalling.