A model for simulating Polygonum aviculare L. seed dormancy loss in relation to stratification temperature was developed. The model employs the lower limit temperature for germination (Tl) as an index of seed population dormancy status. While population mean for Tl (Tl(50)) and Tl distribution within the population (σTl) are allowed to vary as seeds are released from dormancy, other thermal parameters characterizing the germination thermal responses (base, optimal and maximal temperatures, and thermal time required for germination) and the higher limit temperature for germination (Th) are held constant. In order to relate changes in Tl(50) and σTl to variable time and temperature, a stratification thermal time index (Stt) was developed, which consists of the accumulation of thermal time units under a threshold temperature for dormancy loss to occur. Therefore, Tl(50) and σTl varied in relation to the accumulation of Stt according to time and temperature. To derive model equations, changes in seed population thermal parameters were estimated for buried seeds stored at 1.6, 7 and 12°C for 110 d. Seeds were exhumed at regular intervals, and were incubated at 15°C and at a gradually increasing temperature regime in the range 6–25°C. The germination time-course curves obtained were reproduced using a mathematical model. Thermal parameters that best fit simulated and experimentally obtained germination time-course curves were determined. Model performance was evaluated against data of two unrelated experiments, showing acceptable prediction of timing and percentage of germination of seeds exhumed from field and controlled temperature conditions.

Plants undergo a series of physiological, biochemical and molecular changes in response to adverse environmental conditions or stresses such as drought, low temperature or high salt. Several genes and their corresponding proteins have been described that may play a role in withstanding water-deficit-related stresses or full desiccation. In particular, sugars and late-embryogenesis-abundant (LEA) proteins have received the most attention. Plant responses to water-deficit and desiccation have been well-characterized at the molecular level; however, pinpointing the precise roles of the gene products in protecting cells under conditions of water deficit remains a challenging task. While few plants are capable of withstanding full desiccation, most seeds undergo this event as a pre-programmed and final stage in their development. These are the so-called ‘orthodox’ seeds. In contrast to seeds of orthodox species, those of recalcitrant species do not acquire desiccation tolerance during their development and are shed from the parent plant at relatively high water contents. The essential components of desiccation tolerance of seeds are likely to involve the ability to effect repair upon subsequent rehydration as well as the ability to accumulate protective substances that limit the amount of damage which otherwise would be caused by water loss. Studies have begun to examine whether the desiccation sensitivity of recalcitrant seeds is at least partially the result of an insufficient accumulation of LEA-type proteins, or whether other factors (including a lack of protective sugars) are more important. This review assesses some of these studies as well as recent research to understand gene and protein function using transgenic host plant systems.

We studied changes in dormancy (as imposed by the different structures surrounding the embryo, namely, endosperm, pericarp and glumellae) and its relationship with changes in embryonic ABA levels and sensitivity, in developing grains of two commercial barley cultivars: B 1215 and Quilmes Palomar, which have, respectively, a low and high dormancy level at harvest and, consequently, a contrasting sprouting behaviour in rainy years. Dormancy imposed by endosperm plus pericarp was gradually and similarly alleviated throughout development in both cultivars. The presence of the hull (glumellae), in contrast, completely inhibited germination of grains from both cultivars until physiological maturity (PM). From there on, hull-imposed dormancy was removed abruptly in B 1215 grains, while in Q. Palomar ones, it was removed at a much lower rate. This difference determined the contrasting sprouting behaviour of these two cultivars within the ‘time window’ going from PM to crop harvest. Embryonic ABA content and sensitivity were similar in the two cultivars throughout development until PM. From there on, ABA content and sensitivity in B 1215 embryos declined dramatically coinciding with the abrupt termination of hull-imposed dormancy observed in this cultivar. In contrast, ABA levels in Q. Palomar embryos remained high for longer and sensitivity to ABA declined at a much slower rate. This correlation suggests that hull-imposed dormancy in barley might be regulated by embryonic ABA levels and / or sensitivity. Inhibition of GA synthesis with paclobutrazol applied after anthesis lowered the germination capacity of grains from both cultivars without altering that of the naked caryopses, thus further suggesting that hull-imposed dormancy is under hormonal control.

A method is proposed for assessing the maturity and quality of seeds, based on measuring the amount of chlorophyll fluorescence (CF) signals of intact seeds. In general, the amount of chlorophyll is directly related to the degreening process and thus the maturity. Cabbage seeds (Brassica oleracea var. capitata) were separated into three subsamples based on the CF signals of the individual intact seeds. Seeds with the lowest amount of CF had the highest percentage of germination and normal seedlings. In a controlled deterioration test, the subsample with the lowest CF signal had slightly lower germination and normal seedling percentages than the non-treated seeds, whereas the seeds with the highest CF signals had much lower germination and normal seedling percentages. Advantages of the CF method for determining seed maturity and seed quality are its high sensitivity and fully non-destructive nature and the high speed at which the fluorescence is generated and measured.

The effects of preconditioning temperature and preconditioning period on the sensitivity of parasitic weed seeds to the synthetic germination stimulant GR24 were studied under laboratory and field conditions. The temperature during preconditioning of Orobanche cumana and Striga hermonthica seeds strongly affected the responsiveness of the seeds to the applied germination stimulant. Preconditioning at an optimal temperature (21°C for O. cumana and 30°C for S. hermonthica) rapidly released dormancy and increased the sensitivity to GR24 by several orders of magnitude. After reaching maximum sensitivity, prolonged preconditioning rapidly induced secondary dormancy, i.e. decreased sensitivity of O. cumana and S. hermonthica to GR24. The rapid change in sensitivity of preconditioned seeds to germination stimulants during prolonged preconditioning was particularly visible at low concentrations of GR24. GR24 at higher concentrations (0.1 and 1 mg l1) usually induced high germination of both species, regardless of the preconditioning period. The striking similarities between the response of parasitic weed seeds to GR24, described here, and results in the literature on non-parasitic wild plant seeds are discussed. Our results show that parasitic weed seeds are highly sensitive to the germination stimulant for a short period of time only, and then enter into secondary dormancy relatively quickly. The similar germination pattern of S. hermonthica seeds preconditioned for prolonged periods of time under laboratory and field conditions suggests that the mechanism observed is of ecological significance.

Seed longevity of plant species is an important topic in restoration management, and little is known about the effects of environmental conditions on seed survival and longevity under natural conditions. Therefore, the effect of groundwater level on the survival of seeds in the soil seed bank of a natural grassland community was investigated. Large soil cores, mesocosms, were sampled from a grassland site and transferred to two basins under a glass roof. The mesocosms were subjected to different groundwater-level treatments (high and low, respectively 5 and 30 cm below the soil surface) for nearly three years. After that period the soil seed bank of the mesocosms was sampled. In total 15 789 seeds of 38 taxa emerged from the experiment. Significant differences between the number of viable seeds that emerged in the two treatments were found for several species. More seeds of Glyceria fluitans, Cardamine pratensis and Myosotis palustris germinated in the high water-level treatment, whereas fewer seeds of Juncus spp., Cerastium fontanum and Stellaria media were found in this treatment than in the low water-level treatment. The experiment showed that the anoxic conditions prevailing in the high water-level treatment were beneficial to the survival of seeds of species of wet grassland communities. Species of dry grasslands, although represented by only two species, survived better under aerobic conditions.

Saline soils contain multiple types of salt, each of which may exert a different effect on seed germination and seedling growth of plants. The effects of five types of salt on the initial growth of Haloxylon ammodendron, a shrub found on both saline and non-saline areas in deserts of China, were investigated. Seeds were incubated at 20°;C in the dark in a solution (0 to –5.1 MPa) of a salt (NaCl, MgCl2, CaCl2, Na2SO4 or MgSO4) or polyethylene glycol (PEG)-6000, or in a salt (NaCl or MgCl2) or PEG solution containing a low concentration of CaCl2. Seed germination, seedling growth and cation (Na+, Mg2+, Ca2+ and K+) contents of seedlings were examined. Each salt had a different effect on seed germination, seedling growth and influx and outflux of cations in the seedlings. In both NaCl and MgCl2 treatments, the addition of low concentrations of CaCl2 favoured seed germination and seedling growth, and reduced K+ outflux from seedlings, but caused no appreciable decrease in the influx of Na+ or Mg2+ into seedlings. Marked abnormalities in seedlings were found only in treatments with Mg2+ salts, but these effects were completely alleviated by a low concentration of Ca2+ (Ca2+/Mg2+ = 0.012). The different responses of the initial growth in H. ammodendron to different isotonic salt solutions were attributed to differences among salt components in membrane permeability, toxicity and effects on functions of the plasma membrane and/or the cell wall.

Desiccation-tolerant (DT) plant germplasm (i.e. seeds, pollen and spores) survive drying to low moisture contents, when cytoplasm solidifies, forming a glass, and chemical reactions are slowed. DT germplasm may survive for long periods in this state, though inter-specific and intra-specific variation occurs and is not currently explained. Such variability has consequences for agriculture, forestry and biodiversity conservation. Longevity was previously considered in the context of morphological features, cellular constituents or habitat characteristics. We suggest, however, that a biophysical perspective, which considers the molecular organization – or structure – within dried cytoplasm, can provide a more integrated understanding of the fundamental mechanisms that control ageing rates, hence the variation of longevity among species and cell types. Based on biochemical composition and physical–chemical properties of dried materials, we explore three types of the interplay between structural conformations of dried cytoplasm and ageing: (1) cells that lack chlorophyll and contain few storage lipids may exhibit long shelf life, with ageing probably occurring through slow autoxidative processes within the glassy matrix as it relaxes; (2) cells with active chlorophyll may die quickly, possibly because they are prone to oxidative stress promoted by the photosynthetic pigments in the absence of metabolic water and (3) cells that lack chloroplasts but contain high storage lipids may die quickly during storage at −20°C, possibly because lipids crystallize and destabilize the glassy matrix. Understanding the complex variation in structural conformation in space and time may help to design strategies that increase longevity in germplasm with generally poor shelf life.

Changes in tomato (Lycopersicon esculentum Mill.) seed quality were monitored during seed development and maturation in glasshouse experiments in 2 years. The end of the seedfilling period (mass maturity) occurred 35–41 d after anthesis (differing among trusses) in 1989 and 42 d after anthesis in 1990. Seed moisture contents at this developmental stage were 53–72% (wet basis), while the onset of ability to germinate (during 21-d tests at 20°/30°C) and the onset of tolerance to rapid enforced desiccation occurred just before (1990) or just after (1989) mass maturity. In 1989, seed quality was assessed primarily by seedling size in a glasshouse experiment; maximum mean seedling dry weight 25 d after sowing was not achieved until 24–40 d after mass maturity. In 1990, seed quality was assessed primarily by germination following storage; maximum normal germination after 35 d in storage at 40 °C with 14 ± 0.5% moisture content was attained 23 d after mass maturity, but with little difference among seed lots harvested 10 d earlier or up to 30 d later. The results contradict the hypothesis that maximum seed quality is attained at the end of the seed-filling period and that seed viability and vigour begin to decline immediately thereafter.

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

Seed development is a series of events involving cell division, followed by cell differentiation and storage activity In legume cotyledons, cell differentiation starts in certain regions and gradually spreads to other parts, thereby building up developmental gradients The entire process appears to be subject to metabolic control The high hexose state of the premature legume embryo as controlled by seed coat-specific invertases favours cell division Differentiation is initiated when hexose decreases and sucrose increases Seed development occurs in a close interaction with seed metabolism and transport processes Movement of photoassimilates from the sieve tubes to the unloading region of the maternal seed tissue is symplasmic and controlled by plasmodesmal passage Sucrose uptake into Vicia faba cotyledons is mediated by a H+-sucrose symporter located in the outer epidermis which generates transfer cells Formation of the sucrose uptake system is induced during the early to mid-cotyledon stage by tissue contact with the maternal seed coat and is controlled by carbohydrate availability In contrast, a hexose transporter gene is also expressed in epidermal cells covering younger, mitotically active regions of the cotyledons The sucrose uptake system apparently generates the high sucrose state immediately preceding the storage phase Sucrose specifically induces storage-associated differentiation processes indicating a specific sucrose-dependent signalling pathway operating in maturing cotyledons Moreover, the mode of sucrose uptake — apoplasmic movement into the epidermal cells with subsequent symplasmic transfer to the storage parenchyma cells — appears to control coordinated cotyledon development Unlike sucrose, amino acid transport into legume cotyledons is passive during early development but at later stages when large amounts of storage proteins are synthesized an additional active uptake system is established to ensure a sufficient supply

Nitrogenous compounds, such as potassium nitrate, potentiate germination of different species of light-requiring seeds. Using light-induced Arabidopsis thaliana seed germination as a model system, our data suggested that only phytochrome A (phyA)-specific induced germination was affected after the exogenous application of nitrates, different nitric oxide (NO)-donors (such as organic nitrates) or sodium nitroprusside. The stimulative effect was very pronounced. Treated seed samples reached maximal germination after very short periods of red-light irradiation. To a far lesser extent, these substances affected phytochrome B (phyB)-specific induced germination. In phyB-specific induced germination, potassium nitrate was most effective, but germination percentages never exceeded 50%. The least effective was sodium nitroprusside, which practically did not affect phyB-specific induced germination. These results were confirmed using corresponding phytochrome mutants.

The cell cycle is the series of molecular events that allows cells to duplicate and segregate their chromosomes to form new cells. The finding that a protein kinase, the product of the yeast cdc2 gene, was fundamental in the regulation of the G2/M and G1/S transitions, associated with unstable proteins named cyclins, opened a very exciting and dynamic research area. The number of gene products that participate in the development and regulation of the cell cycle may be in the hundreds, and there is a high degree of conservation in protein sequences and regulatory pathways among eukaryotes. Although there are clear differences between plants and animals in cell structure, organization, growth, development and differentiation, the same types of proteins and very similar regulatory pathways seem to exist. Seed germination appears to be an excellent model system for studying the cell cycle in plants. Imbibition will reactivate meristematic cells – most initially with a G1 DNA content – into the cell cycle in preparation for seedling establishment. Early events include a thorough survey of DNA status, since the drying process and seed storage conditions reduce chromosomal integrity. The initiation of cell cycle events leading to G1 and S phases, and of the germination process itself, may depend on a G1 checkpoint control. Most, if not all, cell cycle proteins appear to be already present in unimbibed embryos, although there is evidence of protein turnover in the early hours, suggesting the need for de novo protein synthesis. Regulation also may occur at the level of protein modification, because existing G1, S and G2 cell cycle proteins appear to be activated at precise times during germination. Thus, cell cycle control during seed germination may be exerted at multiple levels; however, knowledge of cell cycle events and their importance for germination is still scarce and fragmentary, and different species may have developed unique control mechanisms, more suited to specific germination characteristics and habitat.

Martin's (1946) seed classification system has 10 types based on embryo and endosperm characteristics and two additional types based on seed size: dwarf (0.3–2.0 mm) and micro ( ≤ 0.2 mm). He listed 17 families and 12 genera (in five other families) as having dwarf seeds. Our recent discovery of morphophysiological dormancy in dwarf seeds of several taxa of Campanulaceae and one of Gentianaceae prompted an evaluation of dwarf seeds. Martin's paper contains 37 families with one to several small (0.3–2.0 mm) seeded species that he did not list as being dwarf. Comparison of Martin's dwarf families and the 37 small-seeded non-dwarf families revealed no consistent differences between the two groups in endosperm texture, seed-coat anatomy, embryo morphology, class of seed dormancy or phylogenetic position. Also, Martin's dwarf seeds include a variety of embryo morphologies. Consequently, we have revised Martin's key to seed types. The dwarf category has been removed and the micro category replaced by ‘undifferentiated’ to reflect the state of the embryo in fresh seeds. Further, the key now includes linear fully developed, linear underdeveloped, spatulate fully developed and spatulate underdeveloped seed types, which Martin illustrated but did not include in his key. In the revised key, all seeds are distinguished on the basis of embryo and endosperm characteristics.

Germination and seedling growth of soybean (Glycine max L.) cv. Williams were examined on paper towels pre-moistened with a range of saline solutions (germination: 0–500 mMolal NaCl; seedling growth: 0–330 mMolal NaCl). The Na+, K+ and Ca2+ concentrations in the embryonic axis immediately before germination and in the seedling 3.5 d after germination were measured. Germination decreased at NaCl concentrations of 330 mMolal (81% germination) and above. At 420 mMolal NaCl, only 40% of seeds germinated, and at 500 mMolal NaCl there was no germination. Seedling growth rate decreased drastically with increasing salinity. At 220 mMolal NaCl, seedling growth rate had declined to 5% of the control, whereas at 330 mMolal NaCl seedling growth was almost zero 3–4 d after germination. Thus, soybean seeds were more tolerant of salinity in the germination than in the seedling phase. The results suggest that the greater tolerance of salinity during the germination phase might, in part, be the result of a lower sensitivity to high tissue Na+ concentrations. Germination (40%) was possible at a tissue Na+ concentration in the embryonic axis of 9.3 mg g FW-1, whereas seedling growth was completely inhibited at a tissue Na+ concentration of 6.1 mg g FW-1. Germination at higher tissue Na+ concentrations was associated with higher K+ and Ca2+ concentrations in the embryo axis, compared with growing seedlings, suggesting that these ions may protect the seeds in the pre-germination phase against salinity.

In the physiological sense, germination begins with seed water uptake and ends with the initiation of elongation by the embryonic axis, usually the radicle. The driving forces and constraints on expansion by the embryo are examined, particularly for seeds in which the embryo is surrounded by endosperm and testa tissues that restrict growth. Models have been developed to predict germination based on thermal time, hydrotime and combined hydrothermal time. These population-based models indicate that the timing of germination is closely tied to physiologically determined temperature and water potential thresholds for radicle emergence which vary among individual seeds in a population. The restraint imposed by tissues surrounding the radicle is a major determinant of the threshold water potential. Enzymatic weakening of these tissues is a key event regulating the timing of radicle emergence. Considerable evidence suggests that endo-β-mannanase is involved in this process in a number of species, although it is doubtful that it is the sole determinant of when radicle emergence occurs. Molecular and biochemical studies are revealing the complexity of events occurring in endosperm and embryo cells associated with the completion of germination. Unique permeability properties and the presence of enzymes associated with pathogen resistance suggest additional functional roles for the tissues enclosing the embryo. The insights gained from physiology and modelling are being extended by the application of molecular techniques to identify and determine the function of genes expressed in association with germination. Single-seed assay methods, in vivo reporters, specific modification of gene expression and mutagenesis will be critical technologies for advancing our understanding of germination

Multispectral imaging is a new technology that is being deployed to assess seed quality parameters. Examples of applications in the detection and identification of fungi on seeds are presented, together with an example of the technology used for maturity determination in sugar beet seed. Results from multispectral imaging are compared with reference methods, and a high correlation is found. Applications of the technique for varietal discrimination and insect damage are also presented. There is a need for non-destructive, reliable and fast techniques, and it is concluded that multispectral imaging has potential for seed quality assessment, in particular for those components associated with surface structure and chemical composition, seed colour, morphology and size.

This review provides a revised and expanded word-formula system of whole-seed primary dormancy classification that integrates the scheme of Nikolaeva with that of Baskin and Baskin. Notable changes include the following. (1) The number of named tiers (layers) in the classification hierarchy is increased from three to seven. (2) Formulae are provided for the known kinds of dormancy. (3) Seven subclasses of class morphological dormancy are designated: ‘dust seeds’ of mycoheterotrophs, holoparasites and autotrophs; diaspores of palms; and seeds with cryptogeal germination are new to the system. (4) Level non-deep physiological dormancy (PD) has been divided into two sublevels, each containing three types, and Type 6 is new to the system. (5) Subclass epicotyl PD with two levels, each with three types, has been added to class PD. (6) Level deep (regular) PD is divided into two types. (7) The simple and complex levels of class morphophysiological dormancy (MPD) have been expanded to 12 subclasses, 24 levels and 16 types. (8) Level non-deep simple epicotyl MPD with four types is added to the system. (9) Level deep simple regular epicotyl MPD is divided into four types. (10) Level deep simple double MPD is divided into two types. (11) Seeds with a water-impermeable seed coat in which the embryo-haustorium grows after germination (Canna) has been added to the class combinational dormancy. The hierarchical division of primary seed dormancy into many distinct categories highlights its great diversity and complexity at the whole-seed level, which can be expressed most accurately by dormancy formulae.

A domestic dog was used in two experiments to elucidate the role of epizoochorous seed dispersal of forest plants. First, the dog was walked through forest vegetation at different times of year and its coat analysed for seeds retained within it. The seed content of the coat was compared to seed frequencies in the vegetation. Secondly, seeds of 11 plant species were placed in different positions on the dog, and their persistence in the coat analysed with respect to distance subsequently travelled. The experiments demonstrate that seeds with morphological adaptations to seed dispersal and small seeds of tall species can be caught effectively by a dog’s coat. The morphologically adapted seeds can be dispersed over large distances as long as the dog moves steadily along a road. The same is true for species with small and smooth seeds if they are deposited on the back of the dog, but not if they are placed on its side. Comparisons of these results with the distribution of forest species in a fragmented landscape, indicated that tall species with small seeds and species with morphological adaptations for epizoochorous dispersal are good at colonizing new forest habitats.

The induction of desiccation tolerance in germinated seeds by incubation in PEG for several days is reported. The induction coincided with an increased sucrose content and the formation of heat-soluble proteins. Germinated seeds can serve as a convenient model system in studies of desiccation tolerance. This finding may have important implications for the agricultural industry.