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In seed formation the role of ethylene has received little attention. The data available on zygotic embryogenesis suggest an association of the ethylene biosynthetic pathway and seed maturation. Over the course of dicot embryogenesis, ACC-oxidase mRNA can be expressed in the cotyledons and embryonic axis. However, as maturation proceeds, cotyledonary ACC-oxidase expression disappears. In some seeds that develop primary dormancy, ethylene synthesis can be among the prerequisites for breaking dormancy. Moreover, the persistence of dormancy may be related to the difficulty of the embryonic axis to produce the necessary ethylene levels or to low tissue sensitivity. The use of inhibitors of ethylene biosynthesis or its action has provided data implicating an ethylene requirement for seed dormancy or germination in some species. However, the role of ethylene in germination remains controversial. Some authors hold that gas production is a consequence of the germination process, while others contend that ethylene production is a requirement for germination. Furthermore, among seeds that require ethylene, some are extremely sensitive to the gas, while others require relatively high levels to trigger germination. Recent studies withXanthium pennsylvanicumseeds suggest that β-cyanoalanine-synthase is involved in ethylene-dependent germination. In addition, regulation of the partitioning ofS-adenosyl-L-methionine (AdoMet) between the ethylene vs polyamine biosynthetic pathways may be a way of controlling germination in some seeds. Such regulation may also apply to the reversal of seed thermoinhibition, which can occur when polyamine synthesis is inhibited, thereby strongly channelling AdoMet towards ethylene. The biological models and approaches that may shed additional light on the role of ethylene during seed germination are presented.

The seed is an important organ of higher plants regarding plant survival and species dispersion. The transition between seed dormancy and germination represents a critical stage in the plant life cycle and it is an important ecological and commercial trait. A dynamic balance of synthesis and catabolism of two antagonistic hormones, abscisic acid (ABA) and giberellins (GAs), controls the equilibrium between seed dormancy and germination. Embryonic ABA plays a central role in induction and maintenance of seed dormancy, and also inhibits the transition from embryonic to germination growth. Therefore, the ABA metabolism must be highly regulated at both temporal and spatial levels during phase of dessication tolerance. On the other hand, the ABA levels do not depend exclusively on the seeds because sometimes it becomes a strong sink and imports it from the roots and rhizosphere through the xylem and/or phloem. All theses events are discussed in depth here. Likewise, the role of some recently characterized genes belonging to seeds of woody species and related to ABA signaling, are also included. Finally, although four possible ABA receptors have been reported, not much is known about how they mediate ABA signalling transduction. However, new publications seem to shown that almost all these receptors lack several properties to consider them as such.

Mannans are hemicellulosic polysaccharides in the plant primary cell wall (CW). Mature seeds, specially their endosperm cells, have CWs rich in mannan-based polymers that confer a strong mechanical resistance for the radicle protrusion upon germination. The rupture of the seed coat and endosperm are two sequential events during the germination of Arabidopsis thaliana. Endo-jS-mannanases (MAN; EC. 3.2

Higher plants have several strategies to perpetuate themselves under adequate ecophysiological conditions. The production of heterogeneous seeds is one such strategy. That is, to ensure the survival of the next generation, an individual plant might produce seeds that are heterogeneous with respect to the extent of dormancy, dispersion and persistence within the seed bank. Heterogeneity can affect not only certain physiological and molecular properties related to seed germination, but also such characteristics as colour, size and shape, parameters commonly used to differentiate morphs within a heterogeneous seed population. In heterogeneous seeds, the above features determine seed behaviour and alter their mechanism of germination. In this work, emphasis is placed on the existence of seed mutants having major alterations in characteristics of the testa and hormonal response. These mutants constitute a valuable tool for elucidating the mechanism of dormancy, germination and perpetuation of seeds. Finally, ontogeny and heterogeneity are reviewed, providing the first data related to the possible hormonal control of heterogeneity in seeds. These results raise the hypothesis that one of the factors triggering differences in germination among heterogeneous seeds may be an alteration in the signalling and action mechanism of ethylene and abscisic acid (ABA).

The softening and degradation of the cell wall (CW), often mannan enriched, is involved in several processes during development of higher plants, such as meristematic growth, fruit ripening, programmed cell death, and endosperm rupture upon germination. Mannans are also the predominant hemicellulosic CW polymers in many genera of green algae. The endosperm CWs of dry seeds often contain mannan polymers, sometimes in the form of galactomannans (Gal-mannans). The endo-β-mannanases (MANs) that catalyse the random hydrolysis of the β-linkage in the mannan backbone are one of the main hydrolytic enzymes involved in the loosening and remodelling of CWs. In germinating seeds, the softening of the endosperm seed CWs facilitates the emergence of the elongating radicle. Hydrolysis and mobilization of endosperm Gal-mannans by MANs also provides a source of nutrients for early seedling growth, since Gal-mannan, besides its structural role, serves as a storage polysaccharide. Therefore, the role of mannans and of their hydrolytic enzymes is decisive in the life cycle of seeds. This review updates and discusses the significance of mannans and MANs in seeds and explores the increasing biotechnological potential of MAN enzymes.

DELAY OF GERMINATION-1 (DOG1), is a master regulator of primary dormancy (PD) that acts in concert with ABA to delay germination. The ABA and DOG1 signaling pathways converge since DOG1 requires protein phosphatase 2C (PP2C) to control PD. DOG1 enhances ABA signaling through its binding to PP2C ABA HYPERSENSITIVE GERMINATION (AHG1/AHG3). DOG1 suppresses the AHG1 action to enhance ABA sensitivity and impose PD. To carry out this suppression, the formation of DOG1-heme complex is essential. The binding of DOG1-AHG1 to DOG1-Heme is an independent processes but essential for DOG1 function. The quantity of active DOG1 in mature and viable seeds is correlated with the extent of PD. Thus, dog1 mutant seeds, which have scarce endogenous ABA and high gibberellin (GAs) content, exhibit a non-dormancy phenotype. Despite being studied extensively in recent years, little is known about the molecular mechanism underlying the transcriptional regulation of DOG1. However, it is well-known that the physiological function of DOG1 is tightly regulated by a complex array of transformations that include alternative splicing, alternative polyadenylation, histone modifications, and a cis-acting antisense non-coding transcript (asDOG1). The DOG1 becomes modified (i.e., inactivated) during seed after-ripening (AR), and its levels in viable seeds do not correlate with germination potential. Interestingly, it was recently found that the transcription factor (TF) bZIP67 binds to the DOG1 promoter. This is required to activate DOG1 expression leading to enhanced seed dormancy. On the other hand, seed development under low-temperature conditions triggers DOG1 expression by increasing the expression and abundance of bZIP67. Together, current data indicate that DOG1 function is not strictly limited to PD process, but that it is also required for other facets of seed maturation, in part by also interfering with the ethylene signaling components. Otherwise, since DOG1 also affects other processes such us flowering and drought tolerance, the approaches to understanding its mechanism of action and control are, at this time, still inconclusive.Plants 2020, 9, 480 2 of 20 (e.g., temperature, O 2 and light) [1][2][3][4]. PD dormancy is a notable agronomic feature. Thus, low levels may produce pre-harvest sprouting, while high levels inhibit the rapid and uniform germination rate. In both cases, there is a reduction of crop production [1,4,5]. PD is established and maintained in the viable dry seed, and throughout several molecular paths (e.g., after-ripening (AR) and exposure to gentle cooling) and can be broken gradually [6,7]. On the other hand, non-dormant seeds of various species have been reported to achieve secondary dormancy (SD) upon exposure to unfavorable conditions for germination [1]. SD occurs essentially after seed dispersal and may be induced by environmental interactions or other special conditions. DELAY OF GERMINATION1 (DOG1), a heme-binding protein, was identified in seeds of A. thaliana by using a biparental recombinant inbred popul...

The transition from the dormant to the non-dormant state of a viable and mature seed can take place at low hydration by exposure to air-dry storage conditions (dry afterripening; AR). The events occurring during this loss of dormancy are of considerable physiological, ecological and agricultural interest. AR may be attributable to increased sensitivity to germination-stimulating factors and a widening of the temperature window for germination. Genetic, –omics and physiological studies on this mode of dormancy breaking provide support for a key role of the balance between gibberellin (GA) and abscisic acid (ABA) metabolism and sensitivity. Recent evidence also supports a possible role for ethylene (ET) in this complex signalling network that is necessary for AR implementation. However, hormone-independent signals, such as reactive oxygen species (ROS), nitrate (NO _{3}^{ - } ) or nicotinamide adenine dinucleotide (NAD+), also appear to be involved. The way in which hormone- and non-hormone-signalling pathways affects each other (cross-talk) is still under study. This review provides updated information on the programmes that overcome seed dormancy. Thus, we have reviewed: (1) the –omic status in dry seeds; (2) the relationship between temperature and nitrate signalling and AR; (3) alterations in ABA/GA synthesis and signalling; (4) the action of hormone molecules other than ABA and GA (i.e. ET, salicylic and jasmonic acids); and (5) participation of reactive oxygen species (ROS), NAD+ and protein carbonylation. Taken together, the acquisition and implementation of dry AR involve a complex signalling network that is difficult to disentangle.