This work presents the results of physiological studies developed to understand modifications linked to the reduction of seed dormancy provoked by domestication processes. The experiments performed compared wild and domesticated Cucurbita subspecies and their hybrids developed by reciprocal crossings. Seeds of two accessions of the wild subspecies presented dormancy, but it was largely reduced in seeds from the domesticated genotype, and partially reverted in hybrids, especially in those obtained when the domesticated genotype was used as the mother plant. In addition, naked embryos of all subspecies did not display dormancy when incubation was performed at 28°C, but embryo germination was progressively reduced only in the wild genotype under decreasing incubation temperatures (22 and 16°C). In the embryos, abscisic acid (ABA) concentrations were similar in both domesticated and wild subspecies, whereas in the seed coat, it was threefold higher in the wild subspecies. The naked embryos from the wild subspecies were far more responsive to ABA than those from the domesticated subspecies. These results indicate that dormancy in the wild subspecies is imposed by the seed coat tissues and that this effect is mediated by their high ABA content and the sensitivity of the embryos to ABA. These physiological aspects were apparently removed by domestication along with the temperature-dependent response for germination.

Published information regarding the effect of storage temperature on dormancy alleviation of sunflower achenes is contradictory and ambiguous. In the present study we explored the effect of temperature during dry storage on dormancy release in two sunflower genotypes, including a commercial hybrid and an inbred line. Dry storage at 25°C consistently accelerated dormancy release of achenes compared with 5°C. This response fits the general pattern reported for dry after-ripening in seeds of many other species. Depending on the genotype and the dormancy factor prevailing, higher temperature alleviated embryo dormancy and coat-imposed dormancy. Hormonal pathways involved in these changes were investigated at the physiological level. In both genotypes, sensitivity to abscisic acid (ABA) was reduced by storage at 25°C. Also, but only in one genotype, storage at 25°C reduced ABA levels upon imbibition and increased the response to a gibberellin (GA) synthesis inhibitor and to applied GA3, compared with storage at 5°C; these results support the idea that temperature affects both ABA and GA metabolism and signalling pathways during after-ripening. This information will be useful to define storage conditions for commercial sunflower achenes, and will also help focus future research on the underlying mechanisms of dormancy release during dry after-ripening in sunflower.

In nature, fruit and seed production in many plants have been shown to be pollen limited. Pollen limitation is demonstrated when open-pollinated plants that are hand-supplemented (Ps) with outcross pollen produce more fruits and/or seeds than open-pollinated controls that are not hand-pollinated (Po). There are three categories of results in such studies: Ps > Po, Ps = Po and Ps < Po, in which case pollen limitation indices are positive, zero and negative, respectively. In an index widely used to calculate pollen limitation, 1 – (Po/Ps), the bounds for Ps ≥ Po are 0 to + 1, whereas the bounds for Ps < Po are 0 to –∞. The first aim of this review was to show how the pollen limitation index can be modified so that the bounds of Ps < Po are 0 and –1, whereupon the index gives equal weight to the best performer (Ps or Po) and worst performer (Ps or Po). In addition to seed quantity, pollen supplementation can affect seed quality, including germinability. Thus, our second aim was to summarize the results of studies that have also tested the effect of pollen limitation on seed germination. In short, the 30 case studies in 15 families, 16 genera and 18 species that we identified show that seed germination percentage increased, was not affected or decreased by pollen supplementation in 12, 11 and seven cases, respectively. The effect of pollen limitation on seed germination, which can be quite large, has not been considered in developing population growth models to determine the effect of pollen limitation on λ.

In seasonal climates, germination timing is mainly controlled by temperature, especially in species with physiological seed dormancy. The germination response to temperature may, however, vary among populations across the distribution range of species. Understanding how populations along climate gradients vary in their sensitivity to temperature is important for determining their vulnerability to climate variability and change. Here, we investigated the germination response of two Erica species with physiological seed dormancy (E. australis and E. umbellata) to changes in temperature throughout the seasons (simulated autumn through to spring) and to the local climate in six localities across a latitudinal gradient in western Iberia. Effects were studied with and without exposing the seeds to a heat shock. The local climate of seed provenance emerged as a key factor in modifying the germination sensitivity to germination temperature and their variation through the seasons. Although each species showed idiosyncratic germination responses to temperature treatments and across the gradient, germination of both species was sensitive to warmer temperatures and to a heat shock. Both showed similar seasonal germination patterns: as we moved from south to north, populations tended to have a larger germination peak in spring, which was greater at colder temperatures. We conclude that rising temperatures associated with climate change will affect these species, particularly at their northern ranges, where many seeds will remain dormant during warmer winters. Arguably, models aiming at assessing climate change impacts in these species need to include such variability across latitude.

Nitrate promotes seed germination at low concentrations in many plant species, and functions as both a nutrient and a signal. As a nutrient, it is assimilated via nitrite to ammonium, which is then incorporated into amino acids. Nitrate reductase (NR) catalyses the reduction of nitrate to nitrite, the committed step in the assimilation. Seed sensitivity to nitrate is affected by other environmental factors, such as light and after-ripening, and by genotypes. Mode of nitrate action in seed germination has been well documented in Arabidopsis thaliana and the hedge mustard Sisymbrium officinale. In these species nitrate promotes seed germination independent of its assimilation by NR, suggesting that it acts as a signal to stimulate germination. In Arabidopsis, maternally applied nitrate affects the degree of primary dormancy in both wild-type and mutants defective in NR. This indicates that nitrate acts not only during germination, but also during seed development to negatively regulate primary dormancy. Functional genomics studies in Arabidopsis have revealed that nitrate elicits downstream events similar to other germination stimulators, such as after-ripening, light and stratification, suggesting that these distinct environmental signals share the same target(s). In Arabidopsis, the NIN-like protein 8 (NLP8) transcription factor, which acts downstream of nitrate signalling, induces nitrate-dependent gene expression. In particular, a gene encoding the abscisic acid (ABA) catabolic enzyme CYP707A2 is directly regulated by NLP8. This regulation triggers a nitrate-induced ABA decrease that permits seed germination. This review article summarizes an update of our current understanding of the regulation of seed dormancy and germination by nitrate.

Differences in fruit morphology among or within species might indicate differences in other regenerative traits, such as seed dormancy and germination. In species with physical dormancy (PY), environmental conditions are suggested to be responsible for dormancy break in field. Seeds of Vachellia caven have PY. This species exhibits two fruit morphs highly represented in Córdoba forests, Argentina: one is dehiscent and the other is indehiscent. In this study we performed a burial experiment with the aim to determine if the differences in V. caven fruit morphology were related to different patterns of PY break of their seeds in field conditions. We related these patterns to (1) environmental conditions that could influence the loss of PY, and (2) histological features of the lens zone. Seeds of both morphs exhibited dormancy break within 14 months of the start of the experiment, but with different patterns. The dehiscent morph showed an abrupt percentage of seeds that broke dormancy 14 months after the beginning of the experiment, probably after undergoing environmental changes similar to those suggested by the two-stage softening model. The indehiscent morph showed a gradual increase in seeds that broke dormancy, not clearly related to any of the environmental variables studied. No differences in seed coat structure of the lens zone were observed between morphs. The existence of both morphs could confer the species with higher possibilities of establishing and coping with environmental heterogeneity. Those characteristics contribute to the understanding of the success of this species in open and disturbed environments.

Seeds often survive in the soil in a dormant state, but their persistence is endangered by micro-organisms that could feed on them. Seed–microbe interactions in the soil are, however, poorly understood. We used dormant caryopses of red rice to study the defence response induced by pronase, a mixture of proteases secreted by Streptomyces griseus, a non-pathogenic bacterium. Pronase was shown to activate the plant immune reaction, indicating that its activity was recognized as a potential microbial attack. The defence reaction included extracellular alkalinization and superoxide production, and the former was necessary to activate the latter, since buffering at pH 6 inhibited the oxidative burst. Alkalinization was sufficient to trigger the oxidative burst, as superoxide production increased when caryopses were incubated in buffered solutions of increasing pH without pronase. Release of proanthocyanidins was observed, with or without pronase. These diverse mechanisms are hypothesized to cooperate in reinforcing seed protection. Finally, time profiles of superoxide production by dormant and non-dormant red rice caryopses during imbibition did not support a relationship between extracellular superoxide and dormancy breaking or germination. Thus, the role of this reactive oxygen species in red rice imbibed caryopses appears to be essentially aimed at defence against attacks by challenging micro-organisms.

The objective of this study was to evaluate the effect of day length after flowering on pollen tube elongation, embryo formation and seed development. The quinoa varieties used in this study were Amarilla de Marangani (valley type) and NL-6 (sea-level type). After sowing, the quinoa plants were cultivated in growth cabinets. From sowing to flowering, plants were exposed to a 15 h day length regime. After flowering, the plants were grown under either a 15 h or 11 h day length regime. The elongation of the pollen tube and the formation of the early embryo were not inhibited in either Amarilla de Marangani or NL-6 under the 11 or 15 h day length regimes. Although growth of the embryo in NL-6 was not inhibited by the 15 h day length regime after flowering, the same was not observed in the case for Amarilla de Marangani. In Amarilla de Marangani, seed diameter at 8 and 14 days after flowering under the 11 h day length regime was larger than that of seeds grown under the 15 h day length regime. Thus, the decrease in the number of seeds in Amarilla de Marangani grown under the 15 h day length regime may be caused by the suspension of embryo growth after fertilization.

Interchangeable effects of temperature, moisture content and oxygen on seed longevity have been mostly examined to estimate seed viability during long-term dry storage, whereas few experiments have studied seed viability under near-natural conditions to evaluate seed persistence in the soil. To this end, we artificially aged seeds of Ranunculus baudotii, a hydrophyte widely distributed in temporary ponds constituting an abundant soil seed bank. Seeds were exposed to controlled ageing at three different relative humidities (RH) under both aerobic and anoxic conditions. Their viability, water content, membrane damage, oxidative stress and anti-oxidant enzymatic defence activity were evaluated. Seed survival was longer at higher relative humidity (97% RH), and lowest at a relative humidity (90% RH) simulating moist but not waterlogged soils. Anoxic conditions showed a protective role on viability at lower moisture contents (70% RH). Seed viability was negatively associated with hydrogen peroxide content and correlated with anti-oxidant enzyme activities, but not with membrane damage. Altogether, these results suggest negative roles for moist soils and anoxia in determining seed persistence in the field, but at higher moisture contents the negative effects of anaerobia diminished. The anti-oxidant system activation, even under unfavourable conditions, might recover seeds once all protective processes can operate, pointing out the plasticity of mechanisms involved in seed loss viability.

Seeds offer a unique perspective from which to view biology. An individual seed is an autonomous biological entity that must rely on its own resources (and resourcefulness) to persist after dispersal and to time its transition to germination and seedling growth to coincide with environmental opportunities for survival. At the same time, seed biology in agriculture and ecology is determined largely by the behaviours of populations of individual seeds. The percentage of seeds in a population that is in a particular state (e.g. dormant, germinated, dead) at a given time is a fundamental metric of seed biology. This duality of individual diversity underlying consistent population-wide behaviour patterns can be described quantitatively using population-based threshold (PBT) models. While conceptually simple, these models are highly flexible and can describe the wide diversity of responses of seed populations to temperature, water potential, hormones, oxygen, light, ageing and combinations of these factors. This seed behaviour is linked to respiratory rates of individual seeds, indicating that basic metabolic processes within seeds vary among individuals in accordance with PBT principles. Looking more broadly across microbial, plant and animal biology, examples of cellular diversity in hormonal sensitivity, gene expression, developmental responses and signalling abound. This variation often is termed ‘noise’, and analysis efforts are focused on extracting mean signals from this variation to understand regulatory pathways. However, extension of the PBT approach to the cellular and molecular levels suggests that population sensitivity distributions and recruitment phenomena may underlie many fundamental biological processes. Thus, concepts and quantitative approaches developed for the analysis of seed populations can be applied across biological scales from molecules to ecosystems to interpret inherent biological variation and provide mechanistic insights into the nature of biological regulatory systems.

Seed predation has been suggested to play a crucial role in plant population dynamics and could act as a strong selective force in the evolution of plant traits such as plant phenology, size and number of flowers. The loss of seeds could be particularly threatening for population maintenance of rare plant species. We investigated the influence of seed predators on plant fitness of the rare monocarpic thistle Cirsium decussatum. We tested the following hypotheses: (1) pre-dispersal seed predators reduce the number of dispersed propagules; (2) seed predators select larger inflorescences as oviposition sites; (3) the size of the capitulum is correlated with seed productivity; and (4) seed predators compete for the oviposition sites. We identified ten insect taxa from different taxonomic groups in the capitula of C. decussatum. Terellia longicauda larvae (Diptera: Tephritidae) were the most common. The presence of the flies’ larvae was positively correlated with the predation level in the capitula. Females of T. longicauda selected larger inflorescences for laying eggs that may increase offspring fitness as larger inflorescences promise more food resources. We could not prove the presence of competition between insect species inhabiting the capitula. Our data suggest, however, that T. longicauda, a specialized seminophagous dipteran, is one of the factors that negatively influence the fitness of threatened C. decussatum, lowering significantly the seed pool. Terellia longicauda potentially acts as a strong selective force in the evolution of C. decussatum inflorescence size.

The proteins produced just prior to maturation desiccation in the developing, orthodox seed, are stored in the desiccated state and recruited as the functional proteome upon imbibition. For the resumption of protein function, these stored proteins must be protected from permanent denaturation while dehydrating, throughout desiccation, and during rehydration. For some forms of damage there is the possibility of repair following imbibition potentially coordinated with de-aggregation into monodispersed polypeptides capable of refolding into a functional configuration. While studying aspects of the natural protection and repair mechanism in seeds, evidence has accrued that those proteins directly involved in translation are particular targets of both protection and protein repair. Such a phenomenon was first described by Rajjou et al. (2008) examining the frequency with which proteins involved in translation were identified as differentially abundant between aged and un-aged Arabidopsis seeds and the translational competence of aged versus un-aged seeds. The inference drawn from these observations was that, of all the stored proteins, it is imperative that those involved in translation endure desiccation, quiescence and rehydration in a functional state if the seed is to survive. Proteins involved in any other process other than translation can be replaced from the stored transcriptome or by de novo transcription but no mRNA is of value without the translational machinery. This has become known as ‘Job's rule’ in honour of the laboratory from which this hypothesis was first put forward (Rajjou et al., 2008). We review in this manuscript the evidence accrued to date on which Job's rule is based.

Abies cephalonica cone and seed morphometric characteristics as well as seed germination behaviour were investigated during an 8-year-long diachronic study (2007–2015). The research was carried out in Parnitha National Park, in the part of this Greek endemic, silver fir population that was spared from an unprecedented wildfire (2007). A statistically significant interannual and among-tree variation of cone traits has been identified, except for cone diameter. Cone length is correlated to the number of seeds per cone, while the percentage of empty seeds has been consistently high, with extreme values of 29.3 and 81.8% in a masting (2015) and lean crop (2009) year, respectively. There is also a considerable proportion of dead (including infected) and non-germinated seeds, and the eventual germinable fraction is well below 25% with spikes in masting years (39.4 and 60.9% in 2010 and 2015, respectively). Untreated seeds have been tested at a wide range of constant and alternating temperatures and germination is completed within 4–5 weeks at T ≥ 15°C and ca 15 weeks at 5–10°C. A chilling pretreatment of 4–8 weeks (at 2–4°C) concludes germination at 15–20°C within 2 weeks. A significant white light requirement has been observed for untreated seeds at 15–20°C, while germination was light indifferent at lower or higher temperatures. Data obtained in this study confirm the present, field seed germination during springtime; we may also predict an earlier germination (late autumn to mid-winter) in forest gaps, under the predicted, warmer conditions in the future.

As a contribution to understanding the world biogeography of seed dormancy in the cosmopolitan genus Ilex, we studied seeds of I. argentina, I. brasiliensis, I. brevicuspis, I. dumosa, I. paraguariensis and I. theezans from the subtropical region of Argentina. We hypothesized that seeds of these species have non-deep simple morphophysiological dormancy (MPD). Effects of temperature, cold stratification and gibberellic acid (GA3) on seed germination and embryo growth were tested. Regardless of incubation temperature, little or no germination occurred for any species until ≥6 weeks. There was an up to 3-fold increase in embryo length to seed length (E:S) ratio before seeds germinated, and embryos grew only during warm-stratifying conditions. Seeds of I. brasiliensis, I. brevicuspis and I. theezans had non-deep simple MPD and germinated to ≥80% after 12, 24 and 16 weeks, respectively. Cold stratification increased germination of I. brasiliensis and I. brevicuspis, and GA3 increased the rate but not final germination percentage of I. brasiliensis and I. theezans. Fresh seeds of I. dumosa required 40 weeks of warm stratification to germinate to 53%, while those after-ripened for 2 months germinated to 81% after 30 weeks; this species has intermediate simple MPD. Seeds of I. argentina and I. paraguariensis germinated to 15 and 21%, respectively, after 40 weeks of warm stratification and did not after-ripen or respond to GA3; these seeds have deep simple MPD. This is the first report of intermediate and deep simple MPD that is broken by warm stratification, thereby increasing our knowledge of seed dormancy in Ilex and in subtropical regions.

The genus Aegilops belongs to the secondary gene pool of wheat and has great importance for wheat cultivar improvement. As a genus with only annual species, regeneration from seeds in Aegilops is crucial. In several species in Aegilops, spikes produce different seed morphs, both in size and germination patterns. However, little is known about the ecology of seed germination, nor about the seed longevity in this genus. Here we investigated the germination phenology of Ae. neglecta under laboratory and field conditions and assessed longevity of different seed morphs of five additional Aegilops species using controlled ageing tests. Large seeds were short-lived and germinated faster than small seeds in most of the species. Field experiments with Ae. neglecta showed that large seeds of the dimorphic pair germinated 3 months after dispersal in contrast to 14 months for smaller seeds. Differences in longevity were detected not only in dimorphic seed pairs, but also among seeds from different positions on the spike. Our results indicate that different longevities in seed morphs of Aegilops may reflect a different soil seed bank persistence, with smaller seeds able to maintain a higher viability after dispersal than larger ones, thereby spreading seedling emergence over two years. Differences of seed germination and longevities between seed morphs in Aegilops may have important implications for ex situ seed conservation and reinforce the hypothesis of a bet-hedging strategy in the germination ecology of this genus.

Graduate training programs often produce technically ‘savvy’ scientists with inadequate non-technical skill sets essential for workplace success. The challenges associated with lack of non-technical competency may be exacerbated in highly specialized fields such as seed science. This brief communication describes recent efforts conducted at the 12th Triennial Conference of the International Society for Seed Science to address non-technical skill development for pre-career professionals. Furthermore, I propose a few adaptable ideas to begin confronting the divide between graduate education and professional development.

Morphophysiological dormancy (MPD) is predominantly found in seeds of temperate regions and is uncommon in arid biomes. MPD has been reported in a number of Hibbertia (Dilleniaceae) species of temperate Australia, and in a single species of the arid zone, H. glaberrima. This study aimed to examine the dormancy and germination ecology of seeds of H. glaberrima. Seeds were subjected to temperature stratification treatments designed to mimic summer and autumn conditions in the Pilbara region of Western Australia. Seed germination and embryo growth were measured. We also tested the interaction between temperature stratification and cycles of drying and wetting designed to mimic sporadic rainfall events. All temperature and moisture treatments were tested in combination (+/–) with the smoke-derived chemical karrikinolide (KAR1). Exposing dormant seeds to temperatures suitable for warm stratification (35°C) for ≥ 8 weeks, followed by incubation at 25°C, resulted in significantly higher germination compared with non-stratified seeds. Exposing seeds to dry/wet cycling in conjunction with temperature stratification did not significantly increase germination. Exposure to KAR1 increased germination under most conditions. Once seeds are shed during October to December, they are exposed to hot and sporadically wet conditions over summer, allowing MPD to be overcome in a proportion of the seed population. Seeds may germinate in autumn (March to April), in conjunction with cooler temperatures. More deeply dormant individuals may require more than one summer to overcome dormancy. Similar to other species occurring in fire-prone ecosystems, fire also plays a crucial role in the germination ecology of H. glaberrima.

Habitat-indifferent species that can grow well in both saline and non-saline soils offer a good opportunity for studying seed dormancy and salt tolerance during germination. Here, we assess interactive effects of maternal habitat and incubation conditions on salt tolerance during germination of the habitat-indifferent halophyte Suaeda vermiculata, a common perennial shrub of the arid Arabian deserts. Seeds collected from saline and non-saline habitats were germinated at six salinity levels (0–500 mM NaCl) and incubated at three temperatures and under two light regimes. Studied germination attributes were final germination, germination rate index (GRI) and germination recovery of non-germinated seeds when transferred to distilled water. The results showed insignificant difference in salt tolerance between seeds of the two habitat types at the lower salinities. At higher salinities (400 and 500 mM NaCl), germination of the saline habitat seeds was almost inhibited, but that of non-saline habitat seeds reached various levels depending on light and temperature. Both higher temperatures and darkness resulted in significant reductions of the final germination at the higher salinities. Seeds of the two habitats that did not germinate in the different treatments recovered their germination when transferred to distilled water, indicating that these seeds maintained their viability in saline solution and were able germinate upon the arrival of suitable conditions. Seeds of the two habitats germinated within 2 days at the lower salinities and GRI decreased with the increase in salinity level. Such reduction was obvious for seeds of the saline habitat. Recovery was faster for seeds of the saline habitats, especially for those incubated at higher salinities and in the dark. The difference in dormancy and germination attributes of seeds of the two habitat types reflects ecological adaptations for survival in salt marshes of arid deserts.

Raphanus raphanistrum (wild radish) is a highly competitive weed in winter crops. Integrated weed management practices and decision-making tools benefit from the ability to predict seed longevity and dormancy status in the soil seed bank, as well as time and density of emergence in the field. We wondered if unique values taken from databases could serve for modelling purposes, whatever the origins of the weed populations. We investigated dormancy and longevity of fruits buried in the soil over a four-year seed burial experiment of two highly contrasted populations that differ by their biogeographical origin (oceanic versus continental climate), their habitat (arable field versus undisturbed) and their fruit size (thick versus thin fruit wall). High viability persisted over two years, followed by a rapid reduction, especially for the ‘continental-undisturbed-thin’ population. Dormancy cycling, with dormancy at its lowest in October, was observed for the ‘oceanic-arable-thick’ population, whilst a slow decrease without clear seasonal pattern was found for the other population. These results indicate different ways of regulating seed persistence in the soil, which might be taken into account when building demographic models. These differences could be due to interactions between fruit wall thickness and the other factors; it is possible that a thicker fruit wall increases longevity by promoting dormancy by physical restriction, but depending on temperature. Thinner fruit walls would make plants display other adaptive strategies to maximize survival time in the soil seed bank. Studies involving more populations and isogenic material are needed to address this specific question.