Birds are important seed dispersers for fleshy fruits through their transportation of ingested seeds. The seeds of many species germinate faster and in greater proportions after passing through a digestive tract, although the effects of this passage vary amongst bird and plant species. Many factors determine the germination success of ingested seeds, such as seed scarification during the digestion process, the fertilizing effect of droppings and the removal of pulp surrounding the seeds. In central Spain, the great bustard (Otis tarda) may act as a disperser of European black nightshade (Solanum nigrum). We analysed the germination success of ingested and non-ingested S. nigrum seeds. The fertilizing effect of bustard droppings and the disinhibition effect of the removal of Solanum pulp on final germination percentage, germination speed and viability were also assessed. Although ingested seeds germinated faster than non-ingested seeds, the former showed a lower germination percentage than the latter: 80–87% versus 99%. Droppings and fruit pulp showed no effect on germination enhancement, except in one aspect: the germination speed of non-ingested seeds decreased when they were sprayed with a fruit extract. We confirm that seeds ingested by great bustards had lower germination success than non-ingested seeds. Although seed ingestion by great bustards reduced seedling emergence, the number of emerged seedlings was still quite large. Thus, great bustards may play a role as a S. nigrum seed dispersal vector.

Physical dormancy (PY) in seeds/fruits, which is caused by the water-impermeable palisade layer, has long been considered a mechanism for synchronizing germination to a favourable time for seedling survival and establishment. Recently, a new hypothesis (crypsis hypothesis) was proposed as the main selective factor for the evolution of PY. However, there are some misconceptions in this hypothesis. Our objective is to critically evaluate the crypsis hypothesis and to point out that there are multiple adaptive roles of PY. The fundamental argument in the crypsis hypothesis, that PY evolved as an escape mechanism from predators, is not valid according to the evolutionary theory of Darwin. According to Darwin's hypothesis, variations (dormancy in our case) within a population occur randomly, i.e. there is no direct function of a variation at the time of its origin. Different selection pressures operating in the environment increase or decrease the fitness of individuals with the variation. Water-gap anatomy in seeds/fruits and phylogenetic relationships of species with PY suggest that PY has evolved several times in angiosperms. Thus, we argue that not only predatory pressure but also several other environmental pressures (e.g. proper timing of germination, ultra-drying of seeds, dispersal and pathogens) were involved in increasing the fitness of species producing seeds with PY. The significance of PY in the survival of the species under the above-mentioned environmental pressures and other misconceptions of the crypsis hypothesis are discussed in detail.

Despite the importance of the palm family, Arecaceae, little has been systematically documented about the seed behaviour of the many species. The post-harvest seed behaviour of Phoenix reclinata, the highly utilized wild date palm species distributed along the eastern seaboard of Africa, is investigated in the present study. While both embryo and endosperm water concentration declined as the seeds of Phoenix reclinata matured, they remained relatively high: this is a characteristic of (but not confined to) non-orthodox seeds. The ultrastructure of embryo cells, and the finding that negligible water uptake was required for the initiation of germination, were in keeping with the possible non-orthodox nature of the seeds. A developmental study revealed that between the acquisition of full germinability and complete pre-shedding maturity, germination performance appeared to be constrained, suggesting the presence of an inhibitor. Pre-treatment by soaking, mechanical or acid scarification had no significant promotory effect on either rate or totality of germination of mature P. reclinata seeds, while use of water transiently at 100°C was highly deleterious. However, germination of partially dehydrated seeds was initiated sooner if they had been soaked or scarified. Mature P. reclinata seeds tolerated dehydration to a mean embryo water concentration of 0.40 g g–1 (dry mass basis; dmb), but at 0.14 g g–1, both rate and totality of germination were adversely affected. However, viability of seeds dehydrated to the mean embryo water concentration 0.40 g g–1 declined during storage for 16 weeks. It is concluded that P. reclinata seeds are non-orthodox, and are best categorized as showing intermediate post-harvest behaviour.

Regardless of whether or not seeds of the geographically-widespread Solidago altissima and S. nemoralis were exposed to light in autumn, those ‘dispersed’ in autumn (15/°C) or winter (5°C) required 12 weeks of light in winter to germinate to ≥80% in darkness in spring (2 weeks at 20/10°C). On the other hand, seeds of the narrow-endemic S. shortii dispersed in autumn and exposed to ≥2 weeks of light in early winter germinated to ≥77% in darkness in spring, and those dispersed in winter and exposed to ≥6 weeks of light germinated to ≥82%. S. altissima and S. nemoralis seeds not exposed to light during any season germinated to only 0–1% in darkness in spring, whereas S. shortii seeds germinated to 45–56%. Seeds of S. altissima and S. nemoralis kept in darkness in autumn and winter needed a 1-day (14-h photoperiod) light exposure in spring to germinate to ≥75% in darkness, whereas those of S. shortii required only one 5-s exposure. Cold-stratified (nondormant) seeds of S. altissima, S. nemoralis and S. shortii exposed to light with a high far-red/red ratio germinated to significantly higher percentages than dark controls and freshly-matured and lab-stored seeds. Results of this study suggest that a soil seed bank of S. shortii should be smaller and be depleted at a faster rate than those of S. altissima and S. nemoralis, and portions of the seeds of the three species can germinate in the far-red-enriched light under plant canopies.

The germination of intact, moist-chilled white spruce (Picea glauca [Moench.] Voss.) seeds and liquid–nitrogen–decoated (LN2), unchilled seeds is significantly more than of intact, unchilled seeds and intact, unchilled seeds exposed to LN2. The testa is largely responsible for imposition of dormancy in these seeds, although the megagametophyte and/or nucellus also play a role. One or both of these tissues undergoes significant weakening during moist chilling. Excised embryos from dormant and nondormant white spruce seeds elongate when placed on solid Murashige and Skoog minimal organics medium supplemented with a carbon source (sucrose, glucose or galactose), but elongate very little on medium without sugar. They are not killed by rapid imbibition on unsupplemented media. Thus, embryos of white spruce do not exhibit innate dormancy, but are dependent on a carbon source for elongation, and have dormancy imposed on them by their surrounding structures.

In higher plants, the seed precursor (ovule primordia) is composed of three parts: funiculus, nucellus and chalaza, generating the latter one (II) or two (OI and II) protective maternal integuments (seed coat, SC). The appearance of a viable seed requires the coordinate growth and development of the preceding three compartments. Integuments are essentials for seed life as they nourish, protect and facilitate seed dispersion. Endosperm and integument growth and development are tightly coupled. Gymnosperm and angiosperm ovules are commonly unitegmic and bitegmic, respectively. Unusually, ategmy and threetegmy (OI, II and aril) also exist. The expression of the INO, ATS and ETT genes, involved in integument development, seems to have demonstrated that the fusion of OI and II leads to the appearance of unitegmy in higher plants. Likewise, INO expression also manifests the conservation of OI during evolution. The molecular control of SC development is constituted by a signalling network with still a multitude of gaps. The fertilization-independent development of the ovule is repressed by the FERTILIZATION INDEPENDENT SEED (FIS), a Polycomb-Repressive-Complex-2 (PRC2). Both endosperm and SC development are tightly linked to PRC2 function. As in many other developmental processes, auxin plays an essential role during ovule and SC development. Auxin transport from the endosperm to the integuments is regulated by AGL62 (AGAMOUS-LIKE 62), the encoding gene of which is specifically expressed in the endosperm to suppress its cellularization. In the absence of AGL62 (i.e. agl62 mutants), auxin remains trapped in the endosperm and the SC fails to develop (i.e. seed abortion). This update shows that auxin biosynthesis, transport and signalling play a predominant role and seem to be absolutely required in the pathway(s) that lead to SC formation, most likely not as a unique hormonal component.

The seeds of Schinus molle are referred to as displaying physical dormancy because of their water-impermeable endocarp. Therefore, this work aimed to evaluate the germination of S. molle seeds as related to environmental conditions, scarification, storage time and seed anatomy. Various experiments were conducted to test the alleviation of dormancy in newly collected and stored seeds. Acid-scarified seeds incubated under continuous light at 25°C showed greatest vigour and germination. The separation of seeds by specific gravity revealed a higher germination percentage for those seeds that sank. In addition, dry storage alleviated dormancy with a remarkable increase in the various germination parameters. Overall, germination traits decreased after prolonged storage, but even after 12 months the means for germination parameters for stored seeds were still higher than those of newly collected ones. S. molle seeds remain attached to parts of the fruit mesocarp and endocarp. The mesocarp contains several layers of parenchyma showing secretory cavities. The endocarp consists of three layers of sclereids surrounding the embryo. Acid scarification strongly changed the structure of the external layers in the mesocarp, digesting parenchyma cells and removing the contents from both parenchyma cells and the secretory cavities; this improved water uptake during imbibition, which occurred only at the carpellary hilar slit. In conclusion, S. molle seeds are positively photoblastic and show physiological dormancy which can be alleviated by acid scarification and dry storage. Seeds can be stored for over 12 months without significant losses in germination parameters compared to newly collected seeds.

The seed coat of Trifolium repens L. was studied with special emphasis on the development of the strophiole, which is the site for water entry during imbibition in leguminous seeds. The epidermal cells of the strophiole are longer than the cells in the remainder of the seed epidermis in the mature ovule. During seed development the median cells of the strophiolar epidermis divide periclinally into an outer layer of palisade cells and an inner layer of isodiametric cells. Prior to maturity a fissure is formed between some of the palisade cells in the centre of the strophiole. It is suggested that tension develops between the palisade cells and the iso-diametrical cells during later maturation stages causing the formation of the fissure which it is believed functions in water uptake. It is indicated that the ‘light line’ is caused by alteration of cellulose microfibrillar orientation in palisade cell walls. It is confirmed that removal of the epicuticular wax from hard seeds by rinsing in absolute alcohol or hexane does not induce water imbibition. Only when seed coats are mechanically abraded do hard seeds germinate.

The involvement of environmental factors in dormancy cycling is well known in temperate annual species, but it is not known how interaction between soil temperatures and humidity can modulate dormancy in perennial tropical species. In this study the effects were evaluated of substrate temperature and humidity on the modulation of the acquisition and overcoming of secondary dormancy in the buried seeds of two endemic Eriocaulaceae species from the rocky fields (campos rupestres) vegetation in south-eastern Brazil. Fresh seeds of Comanthera bisulcata and Syngonanthus verticillatus were buried and subsequently maintained at temperatures of 15, 20, 25 and 30°C, under three substrate humidity levels (boggy, humid and humid/dry). The seeds were exhumed every 3 months and tested for germination (20°C, 12 h photoperiod) and viability (tetrazolium test). The seeds of both species acquired dormancy after burial in all of the treatments. During the experimental period they demonstrated cycles of acquisition and overcoming of dormancy that were most evident in the treatments involving alterations of the substrate humidity (humid/dry regime) that coincided with the environmental conditions found naturally in the region of origin of the species. The seeds gradually lost dormancy during the dry period and re-acquired it when exposed again to humidity; dormancy would once again be overcome during the subsequent dry period. Burial promoted the acquisition of dormancy in C. bisulcata and S. verticillatus seeds; the lowest temperature tested favoured overcoming dormancy; and varying the humidity regime signalled the acquisition and the overcoming of secondary dormancy.

Heretofore, no detailed account was available on seed dormancy and germination of a member of the Colchicaceae (Liliales). Thus, the primary aim of this study was to do an in-depth investigation of the temperature requirements for dormancy break and germination in seeds of Merendera montana (Colchicaceae) at the embryo and whole-seed levels under near-natural temperatures in a non-heated frame shade-house and under controlled conditions in the laboratory. Mean embryo length in fresh seeds was c. 0.57 mm and embryos had to grow to at least 1.30 mm before radicle emergence. Embryos grew to full size and seeds completed germination (radicles emerged) when they were stratified at 28/14°C for 60 d followed by a cool temperature for 60 d and then incubated at a cool temperature for 30 d. The optimum cool stratification temperature for dormancy-break was 10°C. Thus, after the moist pretreatment at 28/14°C+10°C, radicle emergence was>93% at all incubation temperatures (5, 15/4 and 20/7°C). In its natural habitat, M. montana seeds are dispersed in June, the embryo elongates to full size in autumn and radicles emerge from early November to early February. Although the shoot does not emerge until March and April, it is not physiologically dormant. The shoot emerged from 80% of the radicle-emerged seeds in 13 d at 20/7°C without a previous cold pretreatment. Seeds of M. montana have non-deep complex morphophysiological dormancy, C1b1aB-C1a. This is the first study on seeds with complex MPD to show a delay in shoot emergence following root emergence despite the shoot being physiologically non-dormant.

We report preliminary experiments to test the hypothesis that two sugars are involved in the acquisition of desiccation tolerance in immature maize embryos. Sucrose and raffinose were quantified during embryo development, at 16–27 days after pollination (DAP) and during slow dehydration. Embryos were exposed to slow drying either upon excision (18 and 16 DAP) or after an incubation treatment resulting in a decrease in sucrose (18 DAP). The sugars were separated by HPLC with a refractive-index detector; composition was determined by comparing peak areas of interest with those of individual sugar standards. Although further evidence is needed, our results do not appear to support the hypothesis.

Although mature seeds of the monogeneric conifer family Cephalotaxaceae sensu stricto have underdeveloped embryos, no definitive studies have been done to classify dormancy in this family. Our primary purpose was to determine the kind of dormancy in seeds of Cephalotaxus wilsoniana and to put the results into a broad phylogenetic context for gymnosperms. The species is of horticultural and medicinal value, and information is needed on how to propagate it efficiently from seeds. Embryo growth and germination were monitored for seeds at warm, cold and warm plus cold temperatures, and germination was monitored for seeds subjected to: (1) cold →  warm →  cold →  warm; and (2) warm →  cold →  warm →  cold →  warm temperature sequences. The effects of gibberellic acids GA3 and GA4 were tested on radicle emergence in ungerminated seeds and on shoot emergence in root-emerged seeds. Germination was promoted by ≥ 36 weeks of warm stratification followed by ≥ 8 weeks of cold stratification, but only if seeds were returned to high temperatures. The underdeveloped embryo must increase in length by >120% before the radicle emerges. Neither GA3 nor GA4 was effective in promoting radicle emergence; however, both plant growth regulators increased rate (but not percentage) of shoot emergence in root-emerged seeds. We conclude that seeds of C. wilsoniana have the deep simple level of morphophysiological dormancy (MPD), C1b-C3-B1b; thus, warm stratification followed by cold stratification and then warm-temperature incubation are required for germination. In gymnosperms, MPD is known in cycads, Ginkgo and now in three families of conifers.

Germination timing is a key transition of life history. It not only links subsequent life-history traits, such as plant height and flowering time, but also provides a link to the previous generation through the influence of the maternal environment. Environmental factors may mediate these key links, and consequences of this process may influence species regeneration and dispersal. However, little is known about how environmental factors mediate these key links. Here, germination timing under high (natural light) and low light treatments was estimated for 476 angiosperm species of the eastern Tibetan Plateau grasslands. Furthermore, we used standard (std) and phylogenetic (phy) comparative methods to test if germination timing was associated with plant height, flowering time and maternal habitats under both light treatments. Germination timing was positively correlated with plant height only in low light in std-methods. Germination timing was associated with onset of flowering in both light treatments in std-methods, but only in low light when using phy-methods. Germination timing was positively correlated with elevation only in low light when using both comparative methods. Germination timing was correlated with water in maternal habitat only in high light when using both comparative methods. Germination timing was associated with light in maternal habitat in both light treatments in std-methods, but only in high light when using phy-methods. In summary, light-dependent associations of germination timing with subsequent life-history traits and maternal habitats may influence the probability of plant species life-cycle completion and influence distribution and dispersal of plant species in natural plant communities.

Embryogenesis was induced on cotyledons of immature zygotic embryos of soybean (Glycine max (L.) Merrill) placed on solid medium containing 62.5 mm glutamine, soybean seed growth medium salts and vitamins, and 40 mg I−1 2,4-dichlorophenoxyacetic acid (2,4-D) plus 175 mm maltose, or 8 mg I−1 α-naphthaleneacetic acid (NAA) plus 88 mM sucrose. Somatic embryo development was continued in liquid medium containing 0.16 mg I−1 indole-3-butyric acid and 2.64 mg I−1 abscisic acid, glutamine and salts as above, and 88–438 mM sucrose in progressively increasing steps. Germination was on solid half-strength Murashige-Skoog medium. During maturation, somatic embryos mimicked zygotic embryos in colour, protein concentration, water and solute potentials, and respiration. Protein and lipid accumulated to 329 and 86 g kg−1 dry weight in somatic embryos. Fatty acid composition was similar to that of axes of mature seeds. Before desiccation, the water and solute potentials of maturing somatic embryos declined to −1.13 and −1.99 MPa while turgor increased to 0.86 MPa. Concomitantly, a 60% reduction in activity of the cytochrome oxidase pathway of respiration occurred with somatic embryo maturation at 600 g water kg−1 fresh weight. Although small (about 8 mg per embryo), 60% of the somatic embryos formed roots and shoots during germination after maturation without drying and 30% germinated after drying to 60 g water kg−1 fresh weight. In the greenhouse, somatic plantlets grew to mature plants with seeds.

Neem (Azadirachta indica) is a valuable multipurpose tree of tropical arid and semi-arid regions. The use of its seeds is hindered by their short storage longevity. The possible causes of rapid loss of viability were investigated on different seed lots during exposure to 32% and 75% RH at 20°C. Within 6 months the seeds almost lost germinability at 75% RH, whereas at 32% RH viability decreased only slightly. On rehydration, the axis cells from nongerminable seeds had lost turgor, whereas those from viable seeds were turgescent as visualized by low temperature scanning electron microscopy images of fractured axes. Glutathione oxidation status was used to estimate oxidative stress during storage. Oxidative stress was much higher at 75% RH storage than at 32% RH, mainly caused by the rapid loss of reduced glutathione at 75% RH. Oligosaccharides and phospholipids decreased, and free fatty acids increased during storage at the high RH but remained at a constant level at the low RH. However, the degree of fatty acid unsaturation between viable and nonviable seed lots was similar. During the (slow) dehydration of fresh seeds, total glutathione, oligosaccharides and phospholipids accumulated, particularly in the initially more hydrated seeds. We interpret this accumulation as a post-maturation process associated with acquisition of the capability for long-term survival in the dry state. The mass ratio of oligosaccharides to sucrose was 0.19 on average in dehydrated neem seeds. The data suggest that the storage behaviour of neem seeds has features that characterize it as orthodox.

Obtaining corn hybrid seeds (Zea mays L.) with high vigour depends on the parental lines and the direction of the cross, and this relates to seed desiccation tolerance and composition. This research studied reciprocal crosses between pairs of proprietary, elite parent lines (L1 and L5; L2 and L4) producing hybrid seeds with different qualities attempting to correlate vigour with seed composition, focusing on storage proteins, starch and soluble sugar amounts. Four corn hybrid seed lots produced from reciprocal crosses were compared (HS 15 with HS 51, and HS 24 with HS 42) by assessing germination, vigour, and seedling emergence in the field. Seed composition was assessed in mature, dehydrated seeds. Proteins were extracted, quantified, and analysed by electrophoresis and densitometry. Starch amounts were assessed using a kit and soluble sugars were determined using high performance liquid chromatography with pulsed electrochemical detection. The L1 and L2 lineages, used as female parents, provided seeds with lower vigour; however, the quantification of major protein bands, and sucrose, raffinose and stachyose were similar between seed lot pairs. While both total seed protein and starch varied between reciprocal hybrids for one of the two sets of crosses, the amounts of neither correlated with seed vigour. Interestingly, hybrids with low seed vigour (HS 15, HS 24) accumulated greater amounts of fructose relative to their reciprocal; correlation analysis confirmed these results. We demonstrate different effects on seed vigour dependent on the maternal parent in reciprocal crosses producing hybrid corn seeds. We also show that vigour is negatively correlated with seed reducing sugar contents.

Mammal species contribute in different ways to seed dispersal effectiveness, acting as endozoochorous dispersers, scatter-hoarders and seed predators. Seed removal by these functional mammal groups could be affected by anthropogenic drivers, such as grazing management, a common practice in drylands. We evaluated removal of seeds from a native tree species (Prosopis flexuosa) by terrestrial wildlife mammals with different functional roles, on grazed and ungrazed sites and at different times during the fruiting period of Prosopis. We offered Prosopis fruits, each containing 15 seeds, to animals and used camera traps to identify the species removing them. We obtained the number of seeds removed (1 fruit removed = 15 seeds removed) by each animal species and each functional group. Native and domestic mammals removed 65.4% of the total seeds offered; 69.5% of offered seeds were removed from the grazed area and 61% from the ungrazed site. Considering removal times, 64.25% of offered seeds were removed during the beginning of the fruiting period of Prosopis and 67% towards the end of this period. Small mammals acting either as seed predators (Graomys griseoflavus and Akodon dolores) or scatter-hoarders (Microcavia australis) were the functional mammal groups removing the highest amount of seeds. Seed predators removed more seeds from the ungrazed site, whereas the scatter-hoarder did so at the grazed site. In the ungrazed area, it would be important to ensure habitat heterogeneity in order to improve seed removal by functional groups that disperse seeds, such as endozoochorous dispersers and scatter-hoarders.

A recent hypothesis states that the accumulation of amino-carbonyl reaction products of reducing sugars with proteins (Maillard products) is related to the loss of seed vigour and viability during ageing. Since Maillard products are biochemical end products, their accumulation should provide an index of seed deterioration. A simple fluorescence assay has been used to estimate the quantity of Maillard products in aqueous seed extracts. We tested whether an accumulation of fluorescent compounds correlates with the loss of seed viability during ageing. Fluorescence of extracts from lettuce (Lactuca sativa) embryonic axes increased after controlled deterioration, but was not correlated with ageing period. Fluorescence of extracts from whole broccoli (Brassica oleracea var. italica) seeds was either unchanged or declined during controlled deterioration. Extracts from naturally aged whole seeds of carrot (Daucus carota), tomato (Lycopersicon esculentum), and cauliflower (B. oleracea var. botrytis) had higher fluorescence than controls, but there was no consistent increase in fluorescence in proportion to the loss of viability. Extracts of naturally aged onion (Allium cepa) seeds exhibited less fluorescence than extracts from seeds with higher viability. We conclude that the fluorescence assay does not have general utility as an index of seed deterioration.

To colonize newly disturbed areas, weedy species must disperse their seeds in space and/or in time. Cirsium vulgare has poor dispersal in space, but can form a persistent seed bank. Fresh seeds are capable of germinating in either light/dark or constant dark conditions under favourable diurnal temperatures of 25°C: 10°C. Despite this lack of dormancy, a large proportion of seeds do not germinate in the autumn after dispersal. Variation in germination requirements according to seed colour was also examined. Fresh seeds were placed under one of two overwintering treatments at 5°C, alternating light/dark or constant darkness. Some seeds of all colours: white, light, intermediate, dark and black germinated under these conditions. Six months later, ungerminated seeds subjected to the light/dark treatment did not require light for germination when placed under optimal temperatures. However, most seeds that had overwintered in constant darkness required light for germination. In the field, this induced dormancy would prevent seeds from germinating if buried or located in deep shade. Such seeds have the potential of forming a persistent seed bank. The proportion of dormant seeds did not appear to be associated with seed colour. Seeds incubated at 10°C following overwintering under light/dark conditions germinated to a higher percentage than seeds overwintered in darkness, regardless of the light conditions for germination. The results of this study help to explain the contradiction between delayed germination in the field and the lack of seed dormancy.