Flavonoids are secondary metabolites that are present at high levels in most plant seeds and grains. These compounds appear to play vital roles in defence against pathogens and predators and contribute to physiological functions such as seed maturation and dormancy. At the same time, particular subclasses of flavonoids, such as the proanthocyanidins (condensed tannins), negatively impact the use of seeds and grains in animal feed and can add undesirable qualities to food products for human consumption. This article presents an overview of investigations into the physiological and agronomic aspects of seed and grain flavonoids as well as a review of molecular genetic studies, particularly in maize, Arabidopsis and soybean, where mutants deficient in flavonoid biosynthesis provide useful tools for stydying the metabolic machinery underlying the accumulation of these compounds in reproductive structures.

The dormancy pattern of buried seeds of Datura ferox was studied by testing germination responses to red light (R), far-red light (FR) or darkness after exhumation at regular time intervals. The seeds were deeply dormant at the time of burial in winter. After three months (spring), exhumed seeds began to respond to an R pulse. Summer temperatures did not induce secondary dormancy. During winter of the second year, seed germination after an R pulse increased further; however, germination after an FR pulse or in darkness also increased and reached values close to R. Changes in seed germination after an R or an FR pulse and in darkness were studied in a factorial design experiment, combining different depths of burial (0.5, 5 and 10 cm), presence of vegetation (with or without vegetation canopies), and date of exhumation (4 or 7 months after burial). The seeds placed at 0.5 cm depth did not germinate in any light treatment. A very high sensitivity to light, indicated by the promotion of germination by an FR pulse compared to darkness, was exhibited only in seeds buried at 5 cm in a soil without vegetation. A typical R/FR reversible response was observed in seeds exhumed at 5 and 10 cm in almost all situations of burial; except in a soil without vegetation, at 10 cm depth, in late summer when even dark controls showed high germination. It is concluded that buried seeds of D. ferox are released from dormancy in two steps, the first step during the spring of the first year and the second step during the subsequent winter. This process is not obviously controlled by summer temperatures. The acquisition of a very high sensitivity to light depends on vegetation cover and depth of burial, and thus it is predicted to be affected by agricultural practices.

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.

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 chloride salts of Cd or Ni were added to the nutrient solution in which soybean (Glycine max) plants were grown and the response of the plants to these pollutants examined. Both metals markedly reduced plant biomass and seed production. Accumulation was mostly in the roots. Nickel was more mobile than Cd, reaching higher levels in all plant parts, especially seeds. Within the tissues of mature seeds, the highest concentrations of Ni were found in the axis and testa. The highest concentrations of Cd were in the testa and cotyledon, and the lowest in the axis. When expressed on a per seed basis, metal contents of these organs increased with developmental age. Nickel amounts were lower in the pods than the seeds for all growth stages, however there was no significant difference for Cd. Cadmium reduced mature seed mass. This effect was mostly due to decreased yields of lipids, protein and carbohydrates. Although the number of seeds per pod declined as a response to Ni, seed mass was unaffected and there was no apparent effect on storage reserves.

Methanol accumulates in axis tissues of maturing soybean seeds, correlating with preharvest seed deterioration. Accumulation of methanol appears to be associated with the enzymic demethylation of pectin methyl esters by pectinesterase (PE; EC 3.1.1.11). To characterize PE in developing and maturing soybean (Glycine max (L.) Merrill) seeds, enzyme activity was assayed in axis and cotyledon tissues. Activity per g fresh weight was 20–25 times higher in axes than in cotyledons with highest activities between 45 and 60 days after flowering (DAF). Twenty to 33% of the total PE activity was in the ‘soluble’ form (extracted with water, 0.5 M sucrose, 1 M sucrose and water). Soluble and cell-wall-bound PE (subsequently extracted with 1 M NaCI) were purified and characterized from axes of seeds at 45–60 DAF. Purification of PE was achieved through concentration of extracts by ultra-filtration, precipitation with ammonium sulfate (30–80% saturation), dialysis, gel filtration on Sephadex G-75 columns, and ion exchange chromatography on CM Sepharose CL-6B. Further purification of both soluble and bound PE was by isoelectric focusing (IEF) on ultrathin layers of polyacrylamide gel with simultaneous detection of protein and PE activity. It was possible to follow seven bands exhibiting PE activity with pl values between 6.0 and 9.5 in 1 M NaCI-extracts of total homogenates. Differences in the IEF patterns of bound and soluble PE were observed. Whereas the bound enzyme exhibited more basic PE bands (pl 8–9.5), the soluble enzyme had more active bands at pl 6.5, 7.0 and 7.5. The Mr was close to 33 000 and the pH optimum was 7.8 for both soluble and bound PE.

The drying rate of whole seeds of Ekebergia capensis (Meliaceae) was shown to influence the response to desiccation, with rapidly dried seeds surviving to lower water contents. Short-term rapid drying (to water contents higher than those leading to viability loss) actually increased the rate of germination. The form of the time course of decline of axis water content varied with drying rate; slow drying could be described by an exponential function, whereas with rapid drying initial water loss was faster than predicted by an exponential function. These observations suggest that slow drying brought about homogeneous dehydration and that the rapid drying was uneven across the tissue. This raised the possibility that the different responses to dehydration were a function of different distributions of water in the axis tissue under the two drying regimes. However, ultrastructural observations indicated that different deleterious processes may be occurring under the different drying treatments. It was tentatively concluded that a major cause of viability loss in slowly dried material was likely to be a consequence of aqueous-based processes leading to considerable membrane degradation. Uneven distribution of tissue water could not be rejected as a contributory cause of the survival of rapidly dried seeds to low bulk water contents. The differential response to dehydration at different drying rates implies that it is not possible to determine a ‘critical water content’ for viability loss by recalcitrant seeds.

The longevity of neem, Azadirachta indica, seeds from African Sahelian (Burkina Faso) and Asian (Sri Lanka) provenances was studied over two years of storage under different conditions of moisture and temperature. After drying to equilibrium moisture content (MC) at different relative humidities at 20°C, seeds were placed in open storage at 20°C or hermetically sealed in packets at temperatures ranging from −20 to +20°C. There was hardly any difference in storage behaviour between seed batches / lots, whatever their provenance. Seeds originating from mature yellow fruits lived longer than seeds from green or brown fruits. In all storage experiments with seeds having MCs ≥ 10%, viability was preserved best at 10−15°C, indicating that neem seed is chilling (and freezing) sensitive. There was no survival longer than 2 years under these conditions. At MCs of 4–8%, seeds were considerably more tolerant of low temperature storage and had 40–60% viability after 2 years at all temperatures tested (−20 to +20°C). However, the seeds were sensitive to imbibitional stress, which could be alleviated by imbibition at temperatures of 25–30°C or above. The difficult storage behaviour of neem seed seems to stem from: (1) the sensitivity to low temperatures at MCs ≥ 10%; (2) the extreme sensitivity to imbibitional stress after storage at ≤ 8% MC; (3) underestimation of the water activity due to the high oil content of the neem seeds, causing unexpected metabolic stress in the higher MC and temperature range.

The relationship between endosperm cap weakening and endo-β-mannanase activity during priming and the time to germination after priming was studied in tomato (Lycopersicon esculentum) seeds. During priming of seeds in −0.4 MPa PEG, the mechanical restraint of the endosperm cap decreased while the endo-β-mannanase activity in the endosperm cap increased. There was no decrease in required puncture force and no increase in endo-β-mannanase activity in seeds during priming in −1.0 MPa PEG. Two classes of seeds could be distinguished during priming in −0.7 MPa PEG: one with decreased required puncture force and one without. A strong correlation was found between the lowering of the mechanical restraint and endo-β-mannanase activity. It was concluded that individual seeds have to cross a threshold water potential in order to develop enzyme activity and lower their mechanical restraint. A decrease in required puncture force and increase in endo-β-mannanase activity correlated with ice crystal-induced porosity in the endosperm cap cell walls in scanning micrographs. It was presumed that ice crystal-induced porosity reflects cell-wall hydrolysis. Germination time after priming correlated positively with required puncture force during priming, depending on the osmotic potential. Seeds in −1.0 MPa PEG improved their time to germination, without a decrease in the required puncture force. Therefore it was concluded that lowering of the endosperm restraint during priming positively affects the time to germination of primed seeds but is not a prerequisite for rapid germination.

Meadow microcosms were established from seed on low-fertility soil of known seed bank composition, and subjected to manipulations of simulated grazing, cutting date, temperature and fertility for seven years. The composition and density of the seed bank was then determined in five 2-cm soil layers (0–2, 2–4, 4–6, 6–8 and 8–10 cm). The seed bank contained three distinct groups of species: species present in the original soil, sown species, and ‘others’. The seed bank was little affected by the experimental treatments, presumably because the sown species made only a small contribution to the seed bank. Nearly all the species in the original soil are known to possess persistent seed banks and had survived, although at reduced density, for seven years. Density of the most abundant species in this group, Sagina procumbens, had changed very little over seven years, confirming the well-documented longevity of the seeds of this species. Seeds of sown species made up only about a quarter of the seed bank, despite accounting for virtually all the above-ground vegetation. Of the sown meadow species, only Plantago lanceolata and Alopecurus pratensis were relatively abundant in the seed bank. These results strongly support the conclusion of other authors that most meadow species, once lost owing to the effects of fertilizers or inappropriate management, will not reestablish from the seed bank. Among species which were neither sown nor present in the original soil, the majority possessed adaptations for wind dispersal and had presumably dispersed into the experimental plots from outside. The most abundant member of this group, Betula pendula, had dispersed from a nearby tree. Density of Betula seeds declined sharply with depth, consistent with the view that seeds on the soil surface are rapidly lost, mainly through germination, but seeds that become buried survive much better. Seeds of Betula appear to be persistent but not particularly long-lived.

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.

This study examines whether post-dispersal seed predators could be an important selective force in determining the seed bank strategies of grassland plants. It tests the hypothesis that species with persistent seed banks should sustain proportionally less predation of buried seeds than species which have transient seed banks and that this should be true irrespective of seed size. Results are drawn from a field experiment examining the relative susceptibility of surface versus buried seeds for 19 herbaceous taxa exhibiting different degrees of seed bank persistence. The data were consistent with the hypothesis that seed predators (rodents) influence the seed bank characteristics of seeds. Rodents removed proportionally more large seeds than small seeds and removed a smaller proportion of seeds with persistent rather than transient seed banks, independently of seed size. On average, burial reduced seed removal by almost 50%. The decrease in rates of seed removal following burial was marked for seeds with persistent seed banks but negligible for seeds with transient seed banks. Herbaceous plants with relatively large seeds (seed mass > 1 mg) that form persistent seed banks were either completely avoided or only consumed in small quantities by rodents. In contrast, large-seeded species with transient seed banks suffer high rates of seed predation. Models of life-history evolution predict trade-offs between seed dormancy and seed mass since dormancy and seed size are correlated traits that both reduce risk in variable environments and thus will show patterns of negative covariation. This paper presents an alternative explanation for this trade-off based on experimental evidence of a negative relationship between seed bank persistence and predation risk.

Lolium multiflorum Lam., a winter annual grass species originating from the Mediterranean region, has become widespread in native grasslands of the flooding pampa (Argentina). The dynamics of dormancy release of L. multiflorum seeds in simulated natural conditions was studied. Dormancy was overcome by exposing imbibed seeds to mild cool temperatures (15°C or below) for brief periods (pre-chilling). After-ripening progressively released dormancy, at a higher rate in seeds that remained under the canopy (30/20°C) than those that were dry-stored (20°C). Germination was promoted by red light only when the after-ripening requirement was partially fulfilled or after pre-chilling treatment. These results provide useful knowledge for designing grazing strategies aimed at obtaining well-developed ryegrass canopy of high forage quality during winter and spring.