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.

Excised Helianthus annuus L. embryos became dormant during the third week after anthesis. At this stage a short drying treatment (3 d) led to a slight improvement in germination but a 6-week dry storage caused a complete release from dormancy. The short drying treatment, however, elicited the embryos' response to an exogenous concentration of GAs which was unable to promote germination of fresh embryos. It therefore appeared that a short drying treatment changed the sensitivity to GAs but was not capable of directing embryo metabolism completely towards a germinative mode. Moreover, this drying treatment reduced considerably the ABA content in both the axis and the embryo. Nevertheless, no correlation could be established between germinability and the ABA content since the amount of ABA was not modified by the 6-week dry storage. The key step for predisposing the seeds to germinate is the suppression of the capacity for ABA synthesis in the axis, a suppression which takes place during 6-week dry storage.

Seeds of the Brussels sprouts cultivar Asmer Aries and the cauliflower cultivar Hipop were subjected to ageing at 20% moisture content and 45°C for 24 or 30 h, respectively; all seeds retained high germination after ageing. Aerated hydration of unaged and aged seeds of both cultivars for 4–8 h at a range of temperatures (10–30°C), followed by drying, resulted in improved performance, except that germination percentage and rate of cauliflower were lower at 10°C. Thus, all treated seeds showed greater germination rate and seedling root length than the control, which may have resulted from the advancement of the process of germination. The deleterious effect of aerated hydration at 10°C on cauliflower could be explained by damage due to rapid imbibition; seeds that had imbibed slowly to close to full imbibition (41% moisture content) before aerated hydration showed no decrease in germination. The improvement of aged seeds after aerated hydration was also revealed by higher germination after the controlled-deterioration test, which indicated less deterioration in treated seeds. Furthermore, the optimum improvements for all seeds were observed at 25°C and were greater when the water was aerated than non-aerated. These observations indicate the activation of metabolic repair processes during aerated hydration, leading to a reversal of the deterioration sustained during ageing.

Artemisia ordosica, A. arenaria and A. sphaerocephala are semi-shrubs inhabiting desert sand dunes in China and often used to rehabilitate desertified areas. Improvement of dune rehabilitation success by sowing requires better understanding of the processes involved in the control of seed germination and seedling emergence in these species. Thus, (1) effects of temperature, light and osmotica (polyethylene glycol-6000) on seed germination, and (2) effects of seed burial depth in sand and irrigation regime on seedling emergence, were studied under controlled conditions. Seeds of the three species required light for germination, and the light fluence needed for germination was dependent on temperature. Seedling emergence of the three species was maximal (70–94%) for seeds sown at a depth of 2.5 mm, and decreased with increasing seed burial depth when the pots were initially and subsequently treated with 16 mm and 3 mm irrigation at 1-d intervals. However, when the pots were initially and subsequently treated with 8 mm and 3 mm irrigation at 2-d intervals, seedling emergence was almost completely suppressed due to water deficiency in sand. It is suggested that the probability of seed germination and seedling emergence of the three species in the field is very limited, because the light requirement restricts seed germination to shallow sand layers where water is lost rapidly due to evaporation. Temperature appeared to have secondary effects on seed germination, by modifying the light sensitivity of seeds.

The long-term utility of DNA banks is predicated on the stability of DNA during storage. The quality and yield of DNA extracted from seeds from four garden species, which varied in age from 1 to 135 years old, was used to examine the early stages of DNA degradation. Seeds that were 70 years old yielded high molecular weight DNA, which permitted amplification of a 650 bp segment of the internal transcribed spacer (ITS) region of the nuclear ribosomal DNA and a 487 bp segment of the plastid-encoded large subunit of rubisco (rbcL). DNA extracted from seeds estimated to be about 135 years old was degraded, but still permitted amplification when routine extraction procedures were used. The rbcL sequences obtained from c. 135-year-old cucurbit seed DNA were consistent with species identifications based on seed morphological features; however, ITS sequences from the same samples were determined to be of fungal origin. A comparison of our results with studies of DNA stability in leaf specimens suggested that DNA degraded within both tissues, but perhaps more slowly within seeds. Evidence that high-quality DNA can be extracted from old, non-viable seeds expands the utility of seed banks in preserving genetic resources.

Germination and dormancy breaking requirements were studied in Selinum carvifolia (L.) L. and Angelica sylvestris L. (Apiaceae). Seeds of these two species have an underdeveloped embryo and are morpho-physiologically dormant. The embryo does not start to grow until physiological dormancy is broken by cold stratification. Incubating seeds at fluctuating temperatures in the light, after cold stratification, had a stimulating effect on embryo growth and seed germination. Seeds of S. carvifolia and A. sylvestris have non-deep simple morphophysiological dormancy (MPD), since gibberellic acid (GA3) could substitute for cold stratification. This is the first report of non-deep simple MPD that is broken by cold stratification in the Apiaceae. Under natural conditions, physiological dormancy is broken by low temperature conditions during winter. Embryo growth and germination occur in a short time interval when temperatures start rising in early spring. Due to the fact that multiple environmental signals regulate dormancy, seedling emergence in these species is timed very accurately in spring.

Among the forage legumes somatic embryogenesis has been most studied in alfalfa (Medicago sativa L.) reflecting inherent breeding problems and the high demand for seeds of this species. Current methods of somatic embryogenesis are reviewed with emphasis on each of the 3 stages involved: the acquisition of competence, the induction of somatic embryogenesis, and somatic embryo development and maturation. The role of plant growth regulators is particularly stressed. Artificial seed production, the development of reduced-generation synthetics or hybrids and genetic transformation of alfalfa are considered. Research on somatic embryogenesis and plant transformation in a wide range of other forage legumes is examined but it is suggested that the application of artificial seed technology is unlikely in forage legumes other than alfalfa.

Polyclonal antibodies to purified recombinant barley haemoglobin (Hb) have been raised in rabbits and used to investigate its expression in monocotyledonous plants. Very little or no Hb expression was observed in dry barley seeds but germination resulted in the expression of Hb which peaked at 2–3 days after imbibition. Hb expression was also observed in maize, wheat, wild oat and Echinochloa crus-galli seeds during germination. Dissection of tissues from the barley seedlings showed that most of the haemoglobin was expressed in the root and seed coat (aleurone layer), with very little in the coleoptile. Imbibition of half-seeds or excised embryos resulted in the expression of haemoglobin. ATP measurements of barley embryos showed that ATP levels quickly increase after imbibition. α-Amylase activity was also determined in embryos to correlate Hb expression with a well-characterized germination response. The results demonstrate that Hb expression is a normal consequence of germination.

In temperate plant communities, seed size is often inversely related to seed longevity at the species level. We tested if the inverse relationship between seed size and seed longevity holds among individuals within populations of Ambrosia trifida L. (Asteraceae), a summer annual agricultural weed in the eastern United States Corn Belt that typically emerges in two successive flushes: one before 1 May and one after 1 May. The effects of maternal phenotype on seed morphology, seed weight, late emergence (emergence after 1 May) and seed-bank persistence were determined in two 1-year experiments. All seeds were collected from individuals at weekly intervals for 3 weeks. Sixty seeds per collection were used in the analysis. Despite fluctuating environmental conditions during seed development, seed weight within maternal families remained constant for different maturation times. Differences among families constituted 69% of total variation in seed weight, whereas differences within maternal families composed 31% of the variation in seed weight. Percentage late emergence and percentage seed-bank persistence varied among maternal families. In four maternal families, either 0% or 100% of seedlings emerged after 1 May, but a majority of other maternal families produced seedlings both before and after 1 May. Seed-bank persistence rates ranged from 1 to 51% among maternal families. Percentage late emergence and percentage seed-bank persistence were inversely related to maternal-family mean seed weight in 2006, but not in 2005. These results suggest that the inverse relationship between seed size and seed longevity occurs among individuals of a population, but is affected by the environment.

A water relations (hydrotime) model was employed to characterize the responses of lettuce (Lactuca sativa L.) seed germination to temperature, water potential (ψ), endosperm disruption, and the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC). The base water potentials (ψb), standard deviations of base water potential and hydrotime constants (θH) estimated by the model were used to predict the germination rates at reduced ψ and supraoptimal temperatures in the presence or absence of ACC. The distribution of ψb values among seeds in the population (ψb(g)) increased (became more positive) as the imbibition temperature increased, indicating a greater sensitivity of seeds to reduced ψ at thermoinhibitory temperatures. Slitting the seeds at the cotyledonary end to disrupt the integrity of the endosperm extended the high temperature range for germination and maintained lower ψb(g) distributions compared with intact seeds as temperature increased. ACC (10 mm) in the imbibition solution raised the upper temperature limit for germination by about 2°C, and also lowered ψb(g) values by 0.1 to 0.2 MPa. The effect of ACC on ψb(g) was more pronounced at higher imbibition temperatures and in slit seeds. ACC (via conversion to ethylene) extends the high temperature limit for lettuce seed germination by acting in the embryo to maintain a lower ψ threshold for the initiation of growth as temperature increases. Evidence from other species as well suggests that this may be a general mechanism for the action of ethylene in promoting seed germination.

Seed-bank dynamics of C. controversa, a common tree in Japanese temperate forests, were investigated at Ogawa Forest Reserve by direct measurement of seed input, soil seed density, and seedling output (germination of soil seeds), together with two field experiments on postdispersal seed mortality. Soil seed density varied from 4 to 233 m−2 in the site near conspecific adults (<5 m horizontally from crown edge), and 0 to 18 m−2 in far sites (≥5 m) during the 1988–92 period. Seed input occurred only in 1988 and 1990: 190–740 m−2 at the sites near conspecific adults, and 4–20 m−2 at the far sites. Seedling output was observed every year and accounted for 0–15% of the loss from the seed bank, occasionally 30–60%. Seeds buried at 5-cm depth suffered moderate mortality (c. 0.2 year−1) by fungal pathogens. In contrast, seeds experimentally placed on the ground suffered higher mortality (0.5–1 year−1) through predation by ground mammals, probably wood mice. In the experiment, seeds whose mesocarp was removed survived much better. On the other hand, distance itself did not appear to affect post-dispersal mortality of seeds. Seeds in canopy gaps also tended to suffer higher mortality. Current spatial pattern and mode of seed dispersal should decrease the potential ability of the seed bank of this species to colonize safe sites. Rare and unpredictable events might increase the contribution of the seed-bank strategy to regeneration in this forest.

Physical dormancy (PY) occurs in at least 18 angiosperm plant families and is caused by water-impermeable palisade cells in seed (or fruit) coats. Breaking of PY involves disruption or dislodgement of water-gap structures causing the seeds/fruits to become water permeable (non-dormant). The water-gap region is a morphologically distinct area of the seed or fruit coat that forms a water-gap complex. The location, anatomy, morphology and origin of water-gaps can differ between and even within families and genera. Water-gap structures sense environmental conditions that allow seeds with PY to become permeable just prior to the commencement of conditions favourable for germination and plant establishment. There are three basic water-gap morpho-anatomies characterized by the way the water-gap opens: Type-I, Type-II and Type-III. In Type-I water-gaps, specific kinds of cells pull apart to form a surface opening, while in Type-II a portion of the surface structure is pulled away from adjacent cells, opening the water-gap. Type-III is the least common type and has a circular, plug-like structure that is dislodged, whereby water entry occurs. In addition, water-gap complexes are either simple or compound, depending on whether only a single primary water-gap structure is involved in dormancy release or an additional secondary water-gap structure opens, permitting water entry.

Orthodox, desiccation-tolerant seeds lose desiccation tolerance during germination. Here, we quantify the timing of the loss of desiccation tolerance, and explore the implications of this event for seed mortality and the shape of germination progress curves for pioneer tree species. For the nine species studied, all seeds in a seedlot lost desiccation tolerance after the same fixed proportion of their time to germination, and this proportion was fairly constant across the species (0.63–0.70). The loss of desiccation tolerance after a fixed proportion of the time to germination has the implication that the maximum number of seeds in a seedlot that can be killed by a single drying event during germination (Mmax) increases with an increasing time to 50% germination (t50) and an increasing slope of the germination progress curve. Consequently, to prevent the seed population from becoming highly vulnerable to desiccation-induced mortality, species with a greater t50 would be expected to have a shallower germination progress curve. In conclusion, these data suggest that the loss of desiccation tolerance during germination may constitute a significant, but previously unexplored, source of mortality for seeds in seasonal environments with unpredictable rainfall.

A UK seed conservation collection of Anemone nemorosa L. seeds held at the Millennium Seed Bank (MSB) showed low viability in its first post-storage test. Because achenes of A. nemorosa are naturally dispersed when they are green, we tested the hypothesis that seeds may not be fully desiccation tolerant and storable at the time of natural dispersal, and that a post-harvest treatment could increase the proportion of desiccation-tolerant seeds. Achenes harvested at the point of natural dispersal in late May in 2003 and 2004 were either placed immediately on 1% water agar at 20°C (‘laboratory’ treatment), or placed in nylon sachets and buried in leaf litter among plants growing in the wild (‘field’ treatment). Samples were withdrawn at intervals over a period of 168 d and tested for desiccation tolerance (drying to 0.059 g H2O (g DW)− 1) and longevity (controlled ageing at 60% relative humidity and 45°C). An initial increase, followed by a decline, in the proportion of seeds surviving desiccation and in the longevity of both laboratory- and field-treated samples coincided with the development of embryos from globular to heart- and then torpedo-shaped. Developmental arrest was not required for the acquisition of desiccation tolerance, and continued growth and development of the embryo resulted in a loss of desiccation tolerance, similar to that seen in orthodox seeds upon radicle emergence. Furthermore, while A. nemorosa seeds, like many from the Ranunculaceae family, might be described as having morphological or morphophysiological dormancy, this lack of developmental arrest does not fit with the usual concept of dormancy. The implications of these results for the classification systems of seed-storage behaviour and dormancy, and for the long-term conservation of seeds of A. nemorosa, are discussed.

The present study evaluated the size and germination characteristics of seeds of seven species of Syngonanthus (distributed among four sections) that occur in different microhabitats in campos rupestres vegetation in Minas Gerais State, to determine if (1) the germination responses of the various species characterize a single germination pattern for the genus; and (2) there is a correlation between the germination characteristics and the geographic distribution as well as the microhabitat of the different species. The experiments were undertaken with recently collected seeds exposed to a 12-h photoperiod and continuous darkness, at constant temperatures of 10–40°C (at intervals of 5°C) in germination chambers. The results indicated the existence of a pattern of reduced seed size and light requirements for germination within the genus. The geographic distribution of different species of the genus cannot be explained by the responses of seed germination to different temperatures, but suggests some relationship to their edaphic microhabitats. Therefore, germination characteristics of the populations studied may have been selected to colonize specific environments at different soil water conditions.

Seeds have evolved to be highly efficient environmental sensors that respond not only to their prevailing environment, but also their environmental history, to regulate dormancy and the initiation of germination. In the present work we investigate the combined impact of a number of environmental signals (temperature, nitrate, light) during seed development on the mother plant, during post-shedding imbibition and during prolonged post-shedding exposure in both dry and imbibed states, simulating time in the soil seed bank. The differing response to these environments was observed in contrasting winter (Cvi, Ler) and summer (Bur) annual Arabidopsis ecotypes. Results presented show that environmental signals both pre- and post-shedding determine the depth of physiological dormancy and therefore the germination response to the ambient environment. The ecotype differences in seed response to ambient germination conditions are greatly enhanced by seed maturation in different environments. Further variation in response develops following shedding when seeds do not receive the full complement of environmental signals required for germination and enter the soil seed bank in either dry or imbibed states. Species seed dormancy characteristics cannot therefore be easily defined, as seed dormancy is a dynamic state subject to within-species adaptation to local environments.

Historic collections of biological material are important genetic resources for taxonomic, evolutionary and historical research. In this paper we describe a seed collection dating from 1862 to 1918 maintained at the Swedish Museum of Cultural History. The collection contains over 3000 well-documented seed samples of various agricultural crops, mostly cereals. A subset of 100 samples divided over ten species frequently represented in the collection and a range of ages were tested for germinability and DNA preservation. None of these accessions were found to contain viable seeds. DNA extracted from the seeds was degraded, but the amount of degradation varied between species. DNA quality was evaluated by yield, fragment size and size of amplification product. Quality was highest for DNA extracted from Pisum sativum and Vicia sativa. DNA extracted from Brassica napus, Beta vulgaris and Trifolium pratense was more fragmented, and DNA extracted from Triticum aestivum, Secale sereale, Hordeum vulgare, Avena sativa and Phleum pratense was most degraded. Polymerase chain reaction (PCR) amplification of ribosomal DNA fragments of up to 700 bp was permitted for most samples in all species. To test whether single-copy nuclear genes could be amplified from the extracted DNA, microsatellite markers were used on the Pisum sativum and Hordeum vulgare samples. Polymorphisms of microsatellite markers were detected between samples for both species. The results show that the 19th-century seed collection can be utilized to infer genetic relationships among obsolete cultivars as well as for other types of genetic research based on sequence or marker analysis.

Studies were conducted to determine if small embryos (i.e. low embryo length:seed length ratio) in mature dwarf seeds (0.2–2 mm) are underdeveloped. In this case, they would grow (inside the seed) prior to germination, and seeds would have morphological or morphophysiological dormancy. Prior to radicle emergence, embryo length in seeds of Drosera anglica (Droseraceae), Campanula americana, Lobelia appendiculata, L. spicata (Campanulaceae) and Sabatia angularis (Gentianaceae) increased 0, 103, 182, 83 and 57%, respectively. Since embryo growth did not occur in seeds of D. anglica prior to germination, embryos, although small, are fully developed; seeds have only physiological dormancy. The underdeveloped embryo in seeds of C. americana has little or no physiological dormancy; thus, seeds have morphological dormancy. On the other hand, underdeveloped embryos in seeds of L. appendiculata, L. spicata and S. angularis are physiologically dormant, and seeds have morphophysiological dormancy. Therefore, since small embryos in dwarf seeds may or may not be underdeveloped, assignment of seeds to a dormancy class requires that studies be done to determine if embryos grow inside the seed before germination can occur. Such information is important in understanding the evolutionary relationship of the different kinds of seed dormancy.

We tested the germination of fresh and frost-treated seeds of 26 species of southern Mongolian mountain and desert steppes, covering the major growth forms of woody and herbaceous perennials and short-lived species in the region. In the field, germination depends on rains that are largely restricted to the summer months between June and August. Thus, germination tests were performed at alternating conditions of 10°C in darkness and 20°C in light (12 h/12 h), which correspond to temperatures at the study site in early and late summer. Seeds of both woody and herbaceous perennials germinated well under the chosen conditions and apparently did not require stratification or scarification. In contrast, germination of annual species was mostly below 30%, while seed viability was equally high in all three growth forms. Winter conditions, simulated by freezing dried seeds at −18°C, hardly changed seed germination in the perennial species, but several short-lived species responded with increased germination. Short-lived species are not abundant in the real vegetation, which is governed by perennials. Thus, we conclude that the important species in Mongolian mountain steppes germinate readily without a dormancy-breaking treatment. A review of the available literature revealed that a complete lack of dormancy, or presence of only conditional dormancy, is also widely described for other species of Central Asian deserts and steppes, which is in contrast to data from North American prairies.