Exposure to smoke is required for the germination of seeds from dormant genotypes of Nicotiana attenuata, a post-fire annual of the Great Basin Desert. Germination can be elicited by GA1,3,4,7 treatments and inhibited by the GA biosynthesis inhibitor, paclobutrazol (PAC), abscisic acid (ABA) and terpenes leached from unburned litter of the plant’s natural habitat. We analysed the endogenous GA and ABA dynamics during the 22 h after imbibition, when smoke-treated dormant seeds commit to germination. Extractable GA1+3 pools decreased in all seeds, but the decrease was more dramatic within 2 h of smoke exposure, which was followed by an increase between hours 2 and 4. Extractable ABA pools increased shortly after imbibition and remained stable in control, water-treated seeds, but decreased sharply in smoke-treated seeds. PAC completely inhibited smoke-induced germination when seeds were treated for 12 h after smoke exposure, consistent with the requirement of de novo GA synthesis for germination. Smoke treatment in the dark did not result in germination, whereas GA3 treatment did, a result consistent with phytochrome-mediated GA biosynthesis. Smoke exposure dramatically increased the sensitivity of seeds to exogenous GA3 treatments in both the light and dark, and light exposure increased this sensitivity an additional tenfold. Taken together, these results suggest that while germination requires endogenous GA synthesis, the effects of smoke treatment increase GA sensitivity, which is correlated with a decrease in endogenous ABA pools.

Patterns of dormancy and smoke responsiveness in artificially buried seeds were investigated in eight native plant species from the species-rich mediterranean-type climate zone of south-western Australia. A comparison was made between germination and viability behaviour of shelf- and field-soil-stored seed, with and without smoke treatment, at least every 3 months. These comparisons corresponded with each of the four seasons. The species chosen germinated with the aid of smoke under field or glasshouse conditions (termed ‘dormant’; n = 4) or produced low and variable germination under glasshouse conditions with smoke (termed ‘deeply dormant’; n = 4). Three trends were observed in viability of soil-stored seeds over 450 d: no decline, gradual decline or late-onset decline. In addition, various germination responses to soil burial and aerosol smoke were observed. Burial was not required for optimal germination in Anigozanthos manglesii. However, for all other species tested, maximum germination was observed only following a period of burial. This was manifested in a germination response without smoke after a short period of burial (Stylidium affine and Conospermum triplinervium) or a longer period of burial (Conostylis neocymosa, Hibbertia commutata, Leucopogon conostephioides, Stirlingia latifolia and Stylidium crossocephalum). Smoke treatment led to high germination in buried seed of S. affine, S. crossocephalum and H. commutata. The patterns of germination detected in this limited number of species indicate that a variety of mechanisms may exist, both temporally and spatially, in south-western Australian species, and support further research of this type for horticulture and land restoration.

There is a substantial literature on the basic physiology and response to desiccation of recalcitrant seeds, but little is known about their ecology and even less of their evolutionary status. It is difficult to assess the response of early land plants to dehydration, but it is likely that desiccation tolerance in vegetative tissue arose concomitantly with the invasion of the land. Similarly, from the fossil record it is not possible to assess the desiccation response of early seeds, and furthermore, it is difficult to see phylogenetic relationships among species producing recalcitrant seeds. A consideration of the available evidence, however, suggests that the first seeds were desiccation-sensitive, but tolerance evolved early and probably a number of times, independently. The desiccation sensitivity and short life span (generally shorter than the interval between flowering) of recalcitrant seeds have implications in terms of regeneration ecology. A long-term soil seed bank as such does not exist; rather the seeds germinate and form a seedling bank. However, there is a wide range in post-shedding physiology among recalcitrant seed species, and although species producing recalcitrant seeds are common in the humid tropics, they do occur in habitats with more marked seasonal variation. Here regeneration strategies may be more specialized.

Seed dormancy and germination of Senna marilandica and S. obtusifolia were compared in greenhouse and laboratory studies. About 90% of the S. obtusifolia seeds were green and had hard seed coat dormancy, whereas the other 10% were brown and nondormant. Seed-colour morphs did not occur in S. marilandica, and nearly 100% of the seeds had hard seed coat dormancy. Seeds of S. obtusifolia were significantly heavier than those of S. marilandica. Mechanical scarification was very effective in overcoming dormancy in seeds of both species. However, concentrated sulfuric acid, absolute ethanol and boiling water were less effective in breaking dormancy in seeds of S. marilandica than in those of S. obtusifolia. Further, incubating seeds at 30/15 to 40/25°C and dry-heat treatments at 80–100°C were ineffective in breaking dormancy in S. marilandica, but significantly increased germination percentages in S. obtusifolia. In neither species were simulated daily/seasonal temperature shifts effective in breaking dormancy. Scarified seeds of both species germinated over a wide range of temperatures in both light and darkness. Under near-natural temperature conditions, seeds of S. marilandica germinated in spring only, whereas those of S. obtusifolia emerged in late spring and throughout summer. Both species can form a long-lived seed bank. Dormancy break by high field temperatures in seeds of S. obtusifolia allows this species to germinate throughout the warm growing season and thus contributes to its success as a weed in arable crops.

Seeds of legumes are generally considered to have physical dormancy and to be orthodox, but most seed biologists are unaware of the various kinds and combinations of dormancy and storage behaviour in seeds of this family. The aim of our study was to document the dormancy and storage behaviour of seeds of 100 native and introduced tropical Fabaceae species in Sri Lanka and classify them into germination/storage behaviour categories. Moisture content (MC) was < 16% for fresh seeds of 94 species and >29% for those of six. Seeds of these six species had low tolerance for desiccation and for low temperatures. Thus, seeds of six species are non-orthodox and 94 species orthodox. Nine of the 100 species were non-dormant, and 2, 3 and 86 had physiological, physiological epicotyl and physical dormancy, respectively. Six germination/storage behaviour categories were identified among the 100 species. However, as in extratropical regions of the world, orthodox storage behaviour and physical dormancy are characteristic of seeds of the majority of species of Fabaceae in tropical Sri Lanka.

As seed dormancy is released within a seed population, both the rate and percentage of germination increase progressively with increasing dose of a dormancy-breaking treatment or condition. Population-based models can account for this behaviour on the basis of shifting response thresholds as dormancy is alleviated. In particular, hydrothermal time analysis of germination sensitivity to water potential (Ψ) and temperature (T) can describe these features of seed behaviour. We used the hydrothermal time model to analyse the effects of dormancy-breaking treatments on germination of dormant true (botanical) potato (Solanum tuberosum L.) seeds (TPS). After-ripening (37°C and 4% seed moisture content) of TPS for 7 or 30 days partially or fully alleviated primary dormancy. The median base water potential required to prevent germination [Ψb(50)] decreased from –0.25 MPa in control seeds to –0.87 MPa and –1.83 MPa after 7 and 30 days of after-ripening, respectively. In contrast, the base temperature for germination (Tb) was relatively unaffected (0–3.3°C). Fluridone (50 μM), an inhibitor of abscisic acid (ABA) biosynthesis, also promoted germination of dormant TPS and lowered Ψb(50), indicating a role for de novo synthesis of ABA during dormancy maintenance. Moist chilling (3 days at 4°C) or gibberellin (100 μM) alleviated secondary dormancy and lowered Ψb(50) values from –0.08 MPa to –0.36 and –0.87 MPa, respectively. The hydrothermal time model allows quantification of dormancy levels and explains why changes in germination speed and percentage are closely correlated during dormancy alleviation.

Light conditions provide important information about the best time and place for seedling establishment. Photoinhibition of seed germination (PISG), defined as the partial or complete suppression of germination under white light, has been interpreted as a physiological adaptation to avoid germination at or near the soil surface. This review is the first report of an all-inclusive, fully quantitative analysis of PISG in seed plants. Pertinent data available from the published literature for 301 taxa from 59 families and 27 orders were assessed. The association of PISG with several plant and seed traits allowed us to consider the adaptive significance of PISG in relation to plant life histories and the natural environments. As no gymnosperm has been found to be truly photoinhibited, it seems that PISG is apomorphic to flowering plants (especially monocots). Seeds of most taxa with PISG have a dark colour and intermediate mass, mostly in the range 1 to 27 mg. PISG is absent from humid tropical regions and from cold climates, but it is strongly associated with open, disturbed and dry habitats. An intriguing implication of PISG is the formation of a soil-surface seed bank. Taken together, these results clearly indicate that PISG is a physiological adaptation to avoid germination on the soil surface, where conditions are not suitable for seedling establishment. PISG is probably much more frequent in seed plants than previously thought. Thus, laboratory experiments should be conducted under well-characterized light and dark conditions.

To understand the cellular processes related to iron transport and sequestration within the developing pea seed (Pisum sativum), total iron and ferritin iron were analysed in seed coat and embryo tissues of the iron-hyperaccumulating pea mutant, Sparkle [dgl, dgl], and its wild-type parent, cv. Sparkle. For plants grown hydroponically with 2 μM Fe, embryo Fe concentrations averaged 65 μg g−1 dry weight in mature wild-type seeds and 163 μg g−1 dry weight in mature dgl seeds; iron concentrations were also higher in dgl seed coats. Extracted and electrophoretically separated seed proteins were probed with a polyclonal antibody raised against pea seed ferritin. In both genotypes, ferritin was detected in the embryo, but not in the seed coat. Ferritin iron accounted for 92% of the total iron in mature wild-type embryos, but only 42% of the total iron in mature dgl embryos. Radiotracer studies using 59Fe were used to characterize the movement of iron within the seed coat. Unequal distribution of 59Fe in opposing sections taken from the two hemispheres of the seed coat demonstrated that iron was symplastically phloem unloaded. These results suggest that iron resides transiently within the nonvascular seed coat cells and that all cells at the inner surface of the seed coat may be involved in the release of iron to the embryo apoplast. However, the form of iron resident within the seed coat and/or taken up by the embryo is presently unknown.

The relationship between initial mobilization of 11-S globulin storage protein (leading to solubilization of its B-subunit) and germination performance after priming was studied in sugarbeet (Beta vulgaris L.) seeds. Priming was conducted for 2 d either in water (hydropriming) or in –2.0 MPa polyethylene glycol-8000 (osmopriming), at various temperatures ranging from 5 to 40°C and oxygen concentrations in the atmosphere ranging from 0 to 21%. For both types of pre-treatments, the range of temperatures and the concentrations of oxygen which were effective in priming were very similar to those which allowed solubilization of the B-subunit of 11-S globulin, supporting the robustness of this protein marker for optimization of sugarbeet seed priming. Furthermore, the temperature and oxygen dependence of priming efficiency closely paralleled that for germination of the untreated seeds, reinforcing the finding that the beneficial effect of priming corresponded to the advancement of germination sensu stricto (i.e. phase II of the germination process). For priming times longer than 2 d, particularly for osmopriming, there was a very dramatic decrease in germination of the treated seeds. For instance, following a 14 d osmopriming at 25°C as much as 60% of the pre-treated seed population failed to germinate when transferred to water. This loss in germination performance quite closely paralleled degradation of LEA (late embryogenesis abundant) proteins, notably a heat-stable seed-specific protein of about 60 kDa and a seed-specific biotinylated LEA protein.

Seed quality standards enable seed users to achieve their objectives in the establishment of uniform seedlings to a high and reliable level for a range of agricultural and horticultural crops, growing systems and market outlets. Quality standards of commercial seed lots are determined by their positions on the seed survival curves and the shape of their germination progress curves. Although comparative descriptions of germination curves can be achieved by the calculation of the mean germination time (MGT; delay to radicle emergence), single early counts of radicle emergence provide a convenient means of predicting MGT and differences between seed lots. Evidence is presented for an ageing and metabolic repair hypothesis as the overall physiological basis to explain the principles behind the standard germination and vigour tests (ageing, electrolyte leakage, cold test, germination rate and seedling size). The work of the International Seed Testing Association (ISTA) in developing convenient, inexpensive and internationally repeatable tests is illustrated.

A characteristic of recalcitrant seeds is that, if they are maintained under storage conditions that prevent water loss, they will ultimately lose viability. A current view is that hydrated recalcitrant seeds are metabolically active and undergo germination-associated changes in storage. Some of these changes, such as extensive vacuolation and increase in cell size, imply a requirement for water additional to that present in the seed on shedding. It is therefore suggested that, in storage, recalcitrant seeds are exposed to an initially mild, but increasingly severe, water stress. Deleterious events associated with a water stress of considerable duration are suggested to lead ultimately to the death of the tissue.

The damage that occurs on prolonged storage is unlikely to be associated with an inability to form glasses or prevent membrane lipid phase changes, as absolute water contents are higher than those at which these mechanisms become important. It is considered that the most likely process leading to death of water-stressed (as opposed to dehydrated) tissue is a breakdown of co-ordination of metabolism, leading to uncontrolled free-radical-mediated oxidative damage.

It is generally difficult to maintain tissue in a mild water-stressed condition for extended periods. Stored, hydrated, recalcitrant seeds may provide an ideal model system for studying the metabolic effects of prolonged mild water stress.

The effect of rate of dehydration was assessed for embryonic axes from mature seeds of Camellia sinensis and the desiccation sensitivity of axes of different developmental stages was estimated using electrolyte leakage. Rapidly (flash) dried excised axes suffered desiccation damage at lower water contents (0.4 g H2O (g DW)−1) than axes dried more slowly in the whole seed (0.9 g H2O (g DW)−1). It is possible that flash drying of isolated axes imposes a stasis on deteriorative reactions that does not occur during slower dehydration. Differential scanning calorimetry (DSC) of the axes indicated that the enthalpy of the melting and the amount of non-freezable water were similar, irrespective of the drying rate.

Very immature axes that had completed morphogenesis and histodifferentiation only were more sensitive to desiccation (damage at 0.7 g H2O (g DW)−1) than mature axes or axes that were in the growth and reserve accumulation phase (damage at 0.4 g H2O (g DW)−1). As axes developed from maturity to germination, their threshold desiccation sensitivity increased to a higher level (1.3−1.4 g H2O (g DW)−1). For the very immature axes, enthalpy of the melting of tissue water was much lower, and the level of non-freezable water considerably higher, than for any other developmental stage studied.

There were no marked correlations between desiccation sensitivity and thermal properties of water. Desiccation sensitivity appears to be related more to the degree of metabolic activity evidenced by ultrastructural characteristics than to the physical properties of water.

The presence of dehydrins could not be demonstrated in axes of mature, undried recalcitrant seeds of the tropical wetland species Avicennia marina, Barringtonia racemosa, Bruguiera exaristata, Bruguiera cylindrica, Bruguiera gymnorrhiza, Ceriops tagal, Rhizophora apiculata, Rhizophora mucronata and Rhizophora stylosa, but were present in the temperate species Acer saccharinum, Aesculus hippocastanum, Araucaria angustifolia, Camellia sinensis, Castanea sativa, Poncirus trifoliata and Zizania palustris. They were also present in axes of Castanospermum australe (of tropical origin) seeds which underwent development in a temperate climate, and were produced in response to drying in axes of Barringtonia racemosa but not Avicennia marina. The presence of dehydrins was associated with high abscisic acid contents. These proteins may provide protection against low temperatures in temperate seeds and against water loss to which the seeds may be naturally exposed. The presence of dehydrins was unrelated to the evolutionary status of the families studied.

It has been proposed that sucrose and raffinose play a role in the storability of maize seeds. The levels of these sugars in the embryos and the glassy state were compared in maize seeds of contrasting storage stability to determine the relationship between sugar composition, glassy state and the storability of the seed. Sucrose was the predominant sugar but its content was not correlated with good storage. The content of raffinose as a mass fraction of total sugars and the magnitude of the glassy state showed positive correlations with storage stability. It is suggested that in maize seeds dry storage stability is a reflection not of the total soluble sugars, but of the mixture of sucrose with raffinose. Apparently raffinose amplifies the magnitude of the glass signal and this in turn is associated with enhanced storability.

Knowledge about the hormonal control of grain dormancy and dormancy loss is essential in wheat, because low grain dormancy at maturity is associated with the problem of pre-harvest sprouting (PHS) when cool and rainy conditions occur before harvest. Low GA (gibberellin A) hormone sensitivity and high ABA (abscisic acid) sensitivity were associated with higher wheat grain dormancy and PHS tolerance. Grains of two PHS-tolerant cultivars were very dormant at maturity, and insensitive to GA stimulation of germination. More PHS-susceptible cultivars were less sensitive to ABA inhibition of germination, and were either more GA sensitive or germinated efficiently without GA at maturity. As grain dormancy was lost through dry afterripening or cold imbibition, grains first gained GA sensitivity and then lost ABA sensitivity. These changes in GA and ABA sensitivity can serve as landmarks defining stages of dormancy loss that cannot be discerned without hormone treatment. These dormancy stages can be used to compare different cultivars, seed lots and studies. Previous work showed that wheat afterripening is associated with decreasing ABA levels in imbibing seeds. Wheat grain dormancy loss through cold imbibition also led to decreased endogenous ABA levels, suggesting that reduced ABA signalling is a general mechanism triggering dormancy loss.

In the Chihuahuan Desert, there are many cacti species considered to be at risk due to illegal extraction, land-use change and overgrazing. To reduce their illegal extraction, ex situ plant propagation has been suggested. However, the literature regarding seed germination biology of these species is scarce. We investigated the effect of light on germination percentages and germination rate (t50) in seeds of 28 cactus species from the Chihuahuan Desert. Seeds were incubated at a 14-h daily photoperiod (light) and in continuous darkness at 25°C for 30 d, after which seeds failing to germinate in darkness were transferred to light for 30 d. Only 11 of the species had non-dormant seeds, germinating ≥70% in the light; thus an evaluation of the effect of light versus darkness on germination was confined to them. All species were positively photoblastic, and all of them had seeds weighing <1 mg. Ten species did not germinate in darkness, and one species had only 7% germination. From these 11 species, 8 did not germinate to a significantly higher percentage when the same set of seeds was transferred from dark to light, suggesting that darkness had triggered secondary dormancy (skotodormancy). To our knowledge, these results are the first to show that darkness triggers secondary dormancy in cacti. Implications of having a light requirement for germination and having small seeds to accumulate a persistent soil seed bank are discussed. These results contribute to understanding the germination biology of cactus species at risk, and could enhance the propagation of large numbers of cultivated individuals outside their habitats, promoting ex situ conservation.

To elucidate the molecular aspects of cereal endosperm development, a good understanding of the histo-differentiation process is necessary. Reviewing the extensive history of lightand electron-microscopic investigations has revealed similarities between cereals with respect to cell ontogeny. Thus, using barley as a model and taking into account many developmental mutants in this and other species, a model for endosperm differentiation is proposed. Based on this model, molecular studies on endosperm gene-regulation are discussed and the available techniques for further investigations are considered.

After synthesis in the vegetative parts of the plant, assimilates are translocated to fruits through xylem and phloem. Research on factors controlling nutrient transport into developing seeds via the phloem has been stimulated by the development of the empty seed coat technique.

There is a consensus that as assimilates are transported from maternal tissues to filial tissues they are delivered to the extracellular space (the apoplast) separating the two generations, prior to uptake from the apoplast into the tissues of the embryo or endosperm. The empty seed coat technique has been used for the study of several aspects of nutrient transport into seeds, e.g. metabolic control and turgorsensitive transport. The osmotic environment of seed tissues has a strong effect on assimilate transport into empty seeds. Several lines of evidence suggest that one of the main ‘secrets’ of the high sink strength of developing seeds, at least in many taxonomic groups of dicotyledons, is that the sink end of the phloem pathway is ‘bathed’ in an apoplast solution with a high concentration of osmotically active solutes. Data on maize do not fit this pattern. A turgor homeostat mechanism may help to maintain high solute concentrations in the seed apoplast. The apoplast environment of seed tissues may also stimulate synthesis of storage proteins and be involved in the prevention of precocious germination. In addition to the osmotic environment, other factors influencing sink strength are discussed. Some aspects of solute transformation during transport through seed tissues are described.

DNA synthesis following the addition of water to excised embryos of non-dormant rye and to embryos of non-dormant and dormant genetic lines of Avena fatua has been examined. All the samples exhibit an early unscheduled DNA synthesis, have similar responses to DNA polymerase inhibitors and a similar increase in DNA ligase function for the first 24 h of imbibition, irrespective of whether they germinate or remain dormant. A β-polymerase-mediated DNA repair activity is indicated immediately upon imbibition with the stable incorporation of 3H-methyl-thymidine into high molecular weight DNA. Following γ-irradiation of dry or imbibed embryos, inhibitor results suggest the appearance of an additional α- or δ-polymerase activity in the ensuing repair. Abscisic acid permits the early unirradiated repair synthesis, but like aphidicolin, it inhibits replicative DNA synthesis and partly inhibits the post-irradiation incorporation of thymidine. DNA synthesis takes place continuously throughout dormancy of imbibed embryos: precursor incorporation into DNA cannot be chased in the short term and occurs in the absence of an S-phase or endoreduplication of nuclear DNA. It is proposed that dormant imbibed embryos maintain the integrity of the genome by a continuous but slow replacement of DNA steered by a non-amplifying and modified form of replicative DNA synthesis and that abscisic acid may play a determining role in this process.

Intact seeds (seed+endocarp) from freshly harvested fruits of Prunus campanulata were dormant, and required 4–6 weeks of warm followed by 8 weeks of cold stratification for maximum germination percentage. Removing both endocarp and seed coat, however, promoted germination in a high percentage of non-stratified seeds. Treatment of intact, non-stratified seeds with gibberellic acid (GA3) was only partially effective in breaking dormancy. However, GA3 promoted germination of non-stratified seeds in which the endocarp (but not the seed coat) had been removed. The order of abscisic acid (ABA) concentration in fresh seeds was endocarp > seed coat > embryo, and its concentration in endocarp plus seed coat was about 6.2-fold higher than that in the embryo. Total ABA contents of seeds subjected to warm and/or cold moist stratification were reduced 6- to 12-fold. A higher concentration of GA4 was detected in embryos of non-dormant than in those of dormant seeds. Fluridone, a carotenoid biosynthesis inhibitor, was efficient in breaking dormancy of Prunus seeds. Paclobutrazol, a GA biosynthesis inhibitor, completely inhibited seed germination, and the inhibitory effect could be partially reversed by GA4, but not by GA3. Thus, dormancy in P. campanulata seeds is imposed by the covering layers. Dormancy break is accompanied by a decrease in ABA content of the covering layers and germination by an increase of embryonic GA4 content.