The Asteraceae with up to 30,000 species occurs on all continents except Antarctica and in all major vegetation zones on earth. Our primary aim was to consider cypselae dormancy-break and germination of Asteraceae in relation to ecology, vegetation zones and evolution. Cypselae are desiccation-tolerant and in various tribes, genera, species and life forms of Asteraceae are either non-dormant (ND) or have non-deep physiological dormancy (PD) at maturity. All six types of non-deep PD are found among the Asteraceae, and dormancy is broken by cold or warm stratification or by afterripening. Soil cypselae banks may be formed but mostly are short-lived. Much within-species variation in dormancy-break and germination has been found. Using data compiled for 1192 species in 373 genera and 35 tribes of Asteraceae, we considered ND and PD in relation to life form, vegetation zone and tribe. Senecioneae and Astereae had the best representation across the vegetation zones on earth. In evergreen and semi-evergreen rainforests, more species have ND than PD, but in all other vegetation zones, except alpine/high-latitude tundra (where ND and PD are equal), more species have PD than ND. Tribes in the basal and central grades and those in the Heliantheae Alliance have both ND and PD. The high diversity and lability of non-deep PD may have enhanced the rate of species diversification by promoting the survival of new species and/or species in new habitats that became available following globally disruptive events since the origin of the Asteraceae in the Late Cretaceous.

The obligate root parasite Cistanche armena is a recently rediscovered, extremely rare endangered species endemic to Armenia, specifically parasitizing camelthorn (Alhagi maurorum, Fabaceae) and saltwort (Salsola dendroides, Chenopodiaceae). Its populations are reputedly declining due to habitat destruction and biotic impacts. Since the only known means of its reproduction is via the seeds, understanding the mechanisms of breaking C. armena seed dormancy and germination along with the related aspects of the species’ biology is highly important both from fundamental (functional ecology and evolution) and applied (conservation and management) perspectives. Here, we present the first in vitro seed germination protocol for C. armena involving fluridone, a systemic herbicide targeting the carotenoid biosynthetic pathway. In addition, the seed micromorphology of C. armena is described using both light microscopy and lignin autofluorescence visualized by confocal laser scanning microscopy. The actin cytoskeleton in radicle cells of germinated C. armena seedlings is described for the first time, being the proof of seed viability. Further elaboration and application of the proposed germination protocol with the cultivation of C. armena on susceptible hosts are altogether seen as a valuable tool for the conservation of this species.

Reproductive strategies for specific populations are closely related to environmental factors. Consequently, they are fundamental for conservation plans and the management of threatened habitats like salt marshes. From this viewpoint, germination strategy under different temperatures and salt conditions, voltammetric parameters and molecular analysis were performed and compared in six Halocnemum populations (four of H. cruciatum and two of H. strobilaceum) growing on Mediterranean (Balearic, Tyrrenic and Adriatic) and Red Sea coasts to establish the relation to environmental variables. Significant interpopulation differences were found in all the evaluated parameters. The Mediterranean populations showed a variable opportunistic germination strategy that was directly related to the drought period length at the studied sites. Consequently, potential environmental predictors of seed response were identified. The most noteworthy were bioclimate, soil texture, continentality index, winter temperatures and summer precipitations. Additionally, voltammetric parameters were evidenced as indicators of maternal plant stress levels and, thus, as potential determinants of future seed responses. The phylogenetic analyses showed a split into two species that did not correspond to germination response. The phylogeographic analyses showed interpopulation differences in haplotype composition for H. cruciatum, but not for H. strobilaceum. In conclusion, the tight connection between seed responses and the ecological parameters of natural populations as an adaptation for successful seedling emergence was proved regardless of its phylogenetic relations.

Generally, orchids produce dust-like seeds in which endosperm reduction and embryo undifferentiation represent a derived state shared with species in about 11 other plant families. Orchid seeds are proposed to have a special kind of morphological or morphophysiological dormancy. We test this proposition, overcoming several design limitations of earlier studies, specifically that the in vitro germination method for orchid seeds uses pro-oxidants for disinfection and incorporates nitrate in the medium; both ‘treatments’ might contribute to dormancy breaking, potentially confounding judgement on the depth and nature of the dormant state. Seeds of the tropical orchid Dendrobium cruentum Rchb. f., were sown both in vitro, on a nutrient medium, and ex vitro, on plain agar omitting prior disinfection with sodium hypochlorite. Seeds previously stored and fresh seeds were incubated under combinations of vitro conditions, light treatments, constant or alternating temperatures and nitrate concentration. Seeds of D. cruentum are very small but have a large embryo that occupies most of the seed. Over a range of constant temperature seeds germinated to the spherical protocorm stage just as well ex vitro as in vitro. Neither light nor nitrate were prerequisites for ex vitro germination. The ability of D. cruentum seed to germinate in the absence of environmental or chemical stimuli suggests that mature seed can be non-dormant. Our results support the proposition that neither all DUST seed fit a dormancy class nor all orchids produce morphological or morphophysiological seeds. Finally, embryo/seed volume determinations in orchids may prove as valuable in studies on the evolution and ecology of germination and dormancy as embryo:seed ratios in other angiosperm species.

Information in the literature and unpublished results of the authors on Dobinea were used to determine the kind [class(es)] of seed (true seed + endocarp) dormancy and of non-dormancy of genera in all five tribes of Anacardiaceae, and the results were examined in relation to the taxonomic position and endocarp anatomy within the family. Reports of both seed germination and endocarp anatomy were found for 15 genera in tribe Spondiadeae, 6 in tribe Anacardieae, 30 in tribe Rhoeae, 3 in tribe Semecarpeae and 1 in tribe Dobineeae. In Spondiadeae (Spondias-type endocarp), Anacardieae, Semecarpeae and Dobineeae (Anacardium-type endocarp), seeds are either non-dormant (ND) or have physiological dormancy (PD). In Rhoeae (Anacardium-type Rhoeae Groups A, B, C and D endocarps), on the other hand, seeds are ND or have physical dormancy (PY), PD or PY + PD. PY/PY + PD in this tribe seems to be restricted (or nearly so) to Rhus s.s. and closely related genera (e.g. Cotinus, Malosma and Toxicodendron) with an Anacardium-type Rhoeae Group A endocarp. However, seeds of other genera (e.g. Astronium and Schinus) with this type of endocarp and those with Rhoeae Group B (e.g. Pistacia), Group C (e.g. Pentaspadon) and Group D (e.g. Heeria) endocarps are either ND or have PD. The fossil fruit record strongly suggests that present-day relationships between diaspore dormancy (or non-dormancy), endocarp structure and taxonomic position within Anacardiaceae extend back to at least the Palaeogene.

Seed health is an indispensable prerequisite of food security. While the toolkit of plant protection products is currently limited, evidence suggests that the seed microbiome could protect seeds from pathogens. Thus, given their possible disease suppressive potential, we tested 11 different pathosystems to achieve the following proof-of-concept: seed microbiomes can be beneficial for seed health through conferring disease suppression. This study focused on beetroot, onion, spinach, pepper, coriander, red fescue and perennial ryegrass seeds, with each crop being challenged with one or two from a total of six pathogens, namely Pythium ultimum (or a Pythium sp.), Setophoma terrestris, Fusarium oxysporum, Phytophthora capsici, Laetisaria fuciformis and a mix of Puccinia sp. isolates. Each seed lot of each crop was tested with and without treatment with a disinfectant as a proxy for comparing intact seed microbiomes with seed microbiomes after partial elimination by disinfection. We found disease suppression in two pathosystems. Beetroot and spinach seed lots were able to suppress disease caused by P. ultimum when their microbiomes were intact but not after seed disinfection. We speculate that this relates to the microorganisms residing on and in the seed. Yet, seed microbiome disease suppression was not found in all pathosystems, highlighting the variation in seed morphology, plant cultivars, pathogens and seed disinfection treatments. A holistic understanding of the characteristics of seeds that harbour suppressive microbiomes as well as the pathogens that are sensitive to suppression could lead to more targeted and informed seed processing and treatment and, consequently, to the sustainable management of seedling diseases.

Fire stimulates the germination of most seeds in fire-prone vegetation. Fruits of Leucadendron (Proteaceae) are winged achenes or nutlets that correlate with their requirements for smoke and/or heat in promoting germination. We describe five possible smoke–heat dormancy-release/germination syndromes among plants, of which Leucadendron displays three (no response, smoke only, smoke and heat). As seed-coat thickness varies with seed-storage location (plant or soil) and morphology (winged or wingless), we tested its possible role in water uptake and germination. Species with winged seeds achieved 100% germination in the absence of smoke/heat, seed coats were an order of magnitude thinner, and their permeability greatly exceeded that of nutlets. As seed-coat thickness increased (1) imbibitional water uptake declined at a decreasing rate, and (2) the response to smoke, and to a lesser extent heat, increased linearly to reach levels of germination approaching those of winged seeds. For species responsive to smoke and heat, there was no additive effect when applied together, suggesting that they may have promoted the same physiological process. We conclude that seed-coat thickness holds the key to germination requirements in this genus, independent of seed-storage location or morphology. By what mechanisms (1) the smoke response is greater the thicker the seed coat and (2) smoke chemicals might increase water permeability to explain the non-additive effect of smoke and heat, warrant further investigation.

Cumulative oxidative damage from the unavoidable formation of reactive oxygen species (ROS) contributes to seed ageing. Low-molecular-weight (LMW) antioxidants, such as water-soluble glutathione (GSH) and lipid-soluble tocochromanols, can prevent ROS from causing damage, especially when antioxidant enzymes are inactive due to desiccation. However, loss of tocochromanols does not always accompany seed ageing, such as during accelerated ageing or controlled deterioration, despite the presence of oxygen and prevalent loss of GSH. To assess relationships between total germination (TG) and antioxidant changes under storage conditions with practical relevance, commercial seeds of Cucumis sativus, Daucus carota, Helianthus cucumerifolius, Latuca sativa, Lepidium sativum, Phaseolus vulgaris and Raphinus sativus of the same cultivar were obtained over multiple years and stored under ambient conditions (21.9 ± 2.1°C; 36.8 ± 6.6% relative humidity). Sigmoidal fitting of TG revealed time to when 50% of seeds had lost viability, which ranged from <5 years (D. carota) to >15 years (C. sativus). Cellular redox states were quantified via the half-cell reduction potential of LMW thiol/disulphide couples. These negatively correlated with TG (i.e. cell redox states were more oxidized in lots with lower TG), with an average R2 value of 0.62 for the most abundant thiol (GSH, or γ-glutamyl-cysteine in P. vulgaris). Concentrations of tocochromanols positively correlated with TG, with an average R2 value of 0.50 for the most abundant tocochromanol (γ or α in L. sativa and H. cucumerifolius). Therefore, during viability loss under ambient ageing conditions leading to the cytoplasm having a glassy state, the lipid domain in all species experienced oxidative damage.

Oxygen is a major factor of seed germination since it allows resumption of respiration and subsequent metabolism reactivation during seed imbibition, thus leading to the production of reducing power and ATP. Most studies carried out in the 60s to 85s indicate that oxygen requirement depends on the species and is modulated by environmental factors. They have also demonstrated that the covering structures mainly inhibit germination by limiting oxygen supply to the embryo during imbibition through enzymatic oxidation of phenolic compounds by polyphenol oxidases (catechol oxidase and laccase) and peroxidases. Recent use of oxygen-sensitive microsensors has allowed to better characterize the oxygen diffusion in the seed and determine the oxygen content at the level of embryo below the covering structures. Here, I will also highlight the major data obtained over the last 30 years indicating the key role of oxygen in the molecular networks regulating seed germination and dormancy through (1) the hormonal balance (ethylene, ABA and GA), the hormone-signalling pathway and, in particular, the ABA sensitivity, (2) the emerging role of mitochondria in ROS production in hypoxia and (3) the involvement of the N-degron pathway in the turnover of proteins involved in seed tolerance to hypoxia.

Seeds show important variation as plant regenerative units among species, but their evolutionary co-variations with other plant characteristics are still poorly understood. Whilst a positive association of seed mass with genome size (GS) and life forms has already been documented, a broad-scale quantification of their evolutionary correlation and adaptive selection has never been conducted. Here, we tested for correlated evolution of seed mass and GS towards distinct selective regimes related to life form in angiosperms. In particular, we tested the hypothesis that the selection toward lighter seeds and smaller genomes is stronger for annual plants, ensuring high regenerative potential. Using multivariate evolutionary models over a dataset containing 3242 species, we showed an overall positive correlated evolution of seed mass and GS deviating from a pure drift process. Instead, evolutionary changes in seed and genome sizes were driven by adaptive selection towards optimal values differing among life forms. Specifically, the evolutionary optima towards which the seed and genome sizes evolve show a covariation toward small values in annuals, intermediate values in perennial herbs and a trade-off between seed mass and GS in woody plants. Moreover, the evolutionary correlation between seed mass and GS is strongest in annuals as an adaption to complete their life cycle in a short time window, when environmental conditions are favourable for regeneration and development to maturity. The asymmetry in the correlated evolution acting on seed and genome sizes due to life form could explain how life-history traits interplay with functional traits and how plants have evolved diverse successful life-history strategies.

Understanding the relative longevity of different seed lots, perhaps of different species or genotypes, but also following production under different environments or using different cultivation methods, or following different post-harvest treatments, is relevant to anyone concerned with the retention of seed lot viability and vigour during storage. However, different scientists over the years have used different conditions to assess seed lot longevity, as well as different variables as the measure of ‘longevity.’ Here, we give some of the backgrounds to how two standard protocols, with an open and closed system respectively, were derived, and explain why we consider p50, defined as the time during storage when seed lot viability, as measured through a germination test, has declined to 50%, is a suitable longevity trait parameter.

Smoke-derived seed germination is an important trait for plants to colonize postfire habitats. The well-characterized smoke-derived chemicals of karrikins germinate seeds of species not known to occur after fires in nature. Hence, the ecologically relevant germination cues in smoke remain to be explored for native postfire plants. With the fire-chaser, Nicotiana attenuata, we revisit a bioassay-driven fractionation of liquid smoke to identify ecologically relevant germination cues. By combining bioassay-guided fractionation and comparative unbiased metabolomics, we developed a robust and efficient method to identify germination cues in smoke. Syringaldehyde (SAL) was re-identified as a germination cue in fractions of liquid smoke that promote seed germination. SAL was found to be produced during wildfires in the plant's native habitat, efficiently adsorbed to N. attenuata seeds from aqueous solutions and not readily leached from soil and accurately predicted the boundaries of natural fire events that reflect the occurrence of native postfire N. attenuata populations. We propose that SAL is an ecologically relevant germination cue in smoke for this species.

The J. Derek Bewley Career Lectures presented at the triennial meetings of the International Society of Seed Science support early-career seed scientists by providing retrospective views, from those late in their careers, of lessons learned and future implications. Ambition, ability, inspiration, foresight, hard work and opportunity are obvious career requirements. The importance of mentoring and teamwork combined with the clear communication of results, understanding and ideas are emphasized. The role of illustration in research, and its dissemination, is outlined: illustration can support hypothesis development, testing and communication. Climate change may perturb the production of high-quality seed affecting conservation as well as agriculture, horticulture and forestry. An illustrative synthesis of the current understanding of temporal aspects of the effects of seed production environment on seed quality (assessed by subsequent seed storage longevity) is provided for wheat (Triticum aestivum L.) and rice (Oryza sativa L.). Seed science research can contribute to complex global challenges such as future food supplies from seed-propagated crops in our changing climate whilst conserving biological diversity (through seed ecology and technologies such as ex situ plant genetic resources conservation by long-term seed storage in genebanks), but only if that research can be – and then is – applied.

Genetic susceptibility to late maturity alpha-amylase (LMA) in wheat (Triticum aestivum L.) results in increased alpha-amylase activity in mature grain when cool conditions occur during late grain maturation. Farmers are forced to sell wheat grain with elevated alpha-amylase at a discount because it has an increased risk of poor end-product quality. This problem can result from either LMA or preharvest sprouting, grain germination on the mother plant when rain occurs before harvest. Whereas preharvest sprouting is a well-understood problem, little is known about the risk LMA poses to North American wheat crops. To examine this, LMA susceptibility was characterized in a panel of 251 North American hard spring wheat lines, representing ten geographical areas. It appears that there is substantial LMA susceptibility in North American wheat since only 27% of the lines showed reproducible LMA resistance following cold-induction experiments. A preliminary genome-wide association study detected six significant marker-trait associations. LMA in North American wheat may result from genetic mechanisms similar to those previously observed in Australian and International Maize and Wheat Improvement Center (CIMMYT) germplasm since two of the detected QTLs, QLMA.wsu.7B and QLMA.wsu.6B, co-localized with previously reported loci. The Reduced height (Rht) loci also influenced LMA. Elevated alpha-amylase levels were significantly associated with the presence of both wild-type and tall height, rht-B1a and rht-D1a, loci in both cold-treated and untreated samples.

Pre-harvest sprouting (PHS) induced by the absence of seed dormancy causes a severe reduction in crop yield and flour quality. In this study, we isolated and characterized TaABI4, an ABA-responsive transcription factor that participates in regulating seed germination in wheat. Sequence analysis revealed that TaABI4 has three homologues, located on chromosomes 1A/1B/1D. TaABI4 contains a conserved AP2 domain, and AP2-associated, LRP and potential PEST motifs. Putative cis-acting regulatory elements (CE1-like box, W-box, ABRE elements and RY elements) were identified in the TaABI4 promoter region that showed high conservation in 17 wheat cultivars and wheat-related species. Expression profiling of TaABI4 indicated that it is a seed-specific gene accumulating during the middle stages of seed development. Transcript accumulation of TaABI4 in wheat cultivar Chuanmai 32 (CM32, PHS susceptible) was 5.07-fold and 1.39-fold higher than that in synthetic hexaploidy wheat SHW-L1 (PHS resistant) at 15 and 20 DPA, respectively. Six expression quantitative trait loci (eQTL) of TaABI4 on chromosomes 2A, 2D, 3B and 4A were characterized based on the accumulated transcripts of TaABI4 in SHW-L1 and CM32-derived recombinant inbred lines. These QTLs explained 10.7 to 46.1% of the trait variation with 4.53–10.59 of LOD scores, which contain genes that may affect the expression of TaABI4.

A low-cost technique named ‘on-farm’ seed priming is increasingly being recognized as an effective approach to maximize crop establishment. It consists of anaerobically soaking seeds in water before sowing resulting in rapid and uniform germination, and enhanced seedling vigour. The extent of these benefits depends on the soaking time. The current determination of optimal soaking time by germination assays and mini-plot trials is resource-intensive, as it is species/genotype-specific. This study aimed to determine the potential of the seed respiration rate (an indicator of metabolic activity) and seed morphological changes during barley priming as predictors of the priming benefits and, thus, facilitate the determination of optimal soaking times. A series of germination tests revealed that the germination rate is mostly attributable to the rapid hydration of embryo tissues, as the highest gains in the germination rate occurred before the resumption of respiration. Germination uniformity, however, was not significantly improved until seeds were primed for at least 8 h, that is, after a first respiration burst was initiated. The maximum seedling vigour was attained when the priming was stopped just before the beginning of the differentiation of embryonic axes (20 h) after which vigour began to decrease (‘over-priming’). The onset of embryonic axis elongation was preceded by a second respiration burst, which can be used as a marker for priming optimization. Thus, monitoring of seed respiration provides a rapid and inexpensive alternative to the current practice. The method could be carried out by agricultural institutions to provide recommended optimal soaking times for the common barley varieties within a specific region.

Seed conservation of banana crop wild relatives (Musa L. spp.) is limited because of lack of knowledge about their germination ecology. Musa acuminata Colla, the most important banana crop wild relative, is distributed in tropical and subtropical Asian and Pacific rainforests and colonizes disturbed sites. The role of temperature in stimulating/inhibiting germination to detect disturbance when canopy gaps are formed is not well known. We assessed seed germination thermal requirements of three subspecies of M. acuminata using nine seed accessions which had been stored in the Millennium Seed Bank. Diurnally alternating temperature cycles were almost completely essential for germination compared with constant temperatures. Germination was optimal when the upper temperature of a diurnal cycle was at 35°C; the lower temperature of the cycle was less important. Subspecies occurrence coordinates were used to extract climate temperature data which were then compared against the temperature requirements for germination from our experiment results. Maximum temperatures of the warmest month across subspecies ranges were close to but below optimal germination temperatures, as were diurnal ranges, suggesting soil-warming at the micro-climate level following gap creation is important for M. acuminata seed germination. Additionally, pre-treatment for 3 months at 60% relative humidity at constant 25°C improved germination from 14 ± 10 (mean, standard deviation) to 41 ± 29% suggesting a period in the soil seed bank under the canopy may increase sensitivity to alternating temperature cycles. Overall viability was low (49 ± 28%), and considerable variance was caused by the different accessions. Germination remained somewhat inconsistent.

The timing of the germination of seeds is highly responsive to inputs from the environment. Temperature plays a key role in the control of germination, with low temperatures acting to stimulate this developmental transition in many species. In Arabidopsis, extensive gene expression changes have been reported at the whole seed level in response to cold, while much less is known about their spatial distribution across the diverse cell types of the embryo. In this study we examined the spatiotemporal patterns of promoter activity and protein abundance for key gibberellic acid (GA) and abscisic acid (ABA) factors which regulate the decision to germinate both during a time course of germination and in response to cold. Low temperature stimulated the spatial relocalization of these factors to the vasculature. The response of these seeds to dormancy-breaking stratification treatments therefore stimulates the distribution of both positive (GA) and negatively acting (ABA) components to this same cell type. This altered spatial pattern persisted following the transfer of seeds to 22°C, as well as after their rehydration, indicating that this alteration is persistent. These observations suggest that the vasculature plays a role in the low temperature-mediated stimulation of germination in this species, while novel cell types are recruited to promote germination in response to stratification.