Loosening of cell walls is an important developmental process in key stages of the plant life cycle, including seed germination, elongation growth, and fruit ripening. Here, we report direct in vivo evidence for hydroxyl radical ( × OH)-mediated cell wall loosening during plant seed germination and seedling growth. We used electron paramagnetic resonance spectroscopy to show that × OH is generated in the cell wall during radicle elongation and weakening of the endosperm of cress (Lepidium sativum; Brassicaceae) seeds. Endosperm weakening precedes radicle emergence, as demonstrated by direct biomechanical measurements. By 3 H fingerprinting, we showed that wall polysaccharides are oxidized in vivo by the developmentally regulated action of apoplastic × OH in radicles and endosperm caps: the production and action of × OH increased during endosperm weakening and radicle elongation and were inhibited by the germination-inhibiting hormone abscisic acid. Both effects were reversed by gibberellin. Distinct and tissue-specific target sites of × OH attack on polysaccharides were evident. In vivo × OH attack on cell wall polysaccharides were evident not only in germinating seeds but also in elongating maize (Zea mays; Poaceae) seedling coleoptiles. We conclude that plant cell wall loosening by × OH is a controlled action of this type of reactive oxygen species.

The micropylar endosperm cap covering the radicle in the mature seeds of most angiosperms acts as a constraint that regulates seed germination. Here, we report on a comparative seed biology study with the close Brassicaceae relatives Lepidium sativum and Arabidopsis thaliana showing that ethylene biosynthesis and signaling regulate seed germination by a mechanism that requires the coordinated action of the radicle and the endosperm cap. The larger seed size of Lepidium allows direct tissue-specific biomechanical, biochemical, and transcriptome analyses. We show that ethylene promotes endosperm cap weakening of Lepidium and endosperm rupture of both species and that it counteracts the inhibitory action of abscisic acid (ABA) on these two processes. Cross-species microarrays of the Lepidium micropylar endosperm cap and the radicle show that the ethylene-ABA antagonism involves both tissues and has the micropylar endosperm cap as a major target. Ethylene counteracts the ABA-induced inhibition without affecting seed ABA levels. The Arabidopsis loss-of-function mutants ACC oxidase2 (aco2; ethylene biosynthesis) and constitutive triple response1 (ethylene signaling) are impaired in the 1-aminocyclopropane-1-carboxylic acid (ACC)-mediated reversion of the ABA-induced inhibition of seed germination. Ethylene production by the ACC oxidase orthologs Lepidium ACO2 and Arabidopsis ACO2 appears to be a key regulatory step. Endosperm cap weakening and rupture are promoted by ethylene and inhibited by ABA to regulate germination in a process conserved across the Brassicaceae.

Appropriate responses of seeds and fruits to environmental factors are key traits that control the establishment of a species in a particular ecosystem. Adaptation of germination to abiotic stresses and changing environmental conditions is decisive for fitness and survival of a species. Two opposing forces provide the basic physiological mechanism for the control of seed germination: the increasing growth potential of the embryo and the restraint weakening of the various covering layers (seed envelopes), including the endosperm which is present to a various extent in the mature seeds of most angiosperms. Gibberellins (GA), abscisic acid (ABA) and ethylene signaling and metabolism mediate environmental cues and in turn influence developmental processes like seed germination. Cross-species work has demonstrated that GA, ABA and ethylene interact during the regulation of endosperm weakening, which is at least partly based on evolutionarily conserved mechanisms. We summarize the recent progress made in unraveling how ethylene promotes germination and acts as an antagonist of ABA. Far less is known about jasmonates in seeds for which we summarize the current knowledge about their role in seeds. While it seems very clear that jasmonates inhibit germination, the results obtained so far are partly contradictory and depend on future research to reach final conclusions on the mode of jasmonate action during seed germination. Understanding the mechanisms underlying the control of seed germination and its hormonal regulation is not only of academic interest, but is also the ultimate basis for further improving crop establishment and yield, and is therefore of common importance.

SummaryThe evolution of the seed represents a remarkable life-history transition for photosynthetic organisms. Here, we review the recent literature and historical understanding of how and why seeds evolved. Answering the 'how' question involves a detailed understanding of the developmental morphology and anatomy of seeds, as well as the genetic programs that determine seed size. We complement this with a special emphasis on the evolution of dormancy, the characteristic of seeds that allows for long 'distance' time travel. Answering the 'why' question involves proposed hypotheses of how natural selection has operated to favor the seed life-history phenomenon. The recent flurry of research describing the comparative biology of seeds is discussed. The review will be divided into sections dealing with: (1) the development and anatomy of seeds; (2) the endosperm; (3) dormancy; (4) early seed-like structures and the transition to seeds; and (5) the evolution of seed size (mass). In many cases, a special distinction is made between angiosperm and gymnosperm seeds. Finally, we make some recommendations for future research in seed biology.