Regulation of seed dormancy by the maternal environment is instrumental for maximizing plant fitness in Polygonum aviculare.

Emergence at an appropriate time and place is critical for maximizing plant fitness and hence sophisticated mechanisms such as seed dormancy have evolved. Although maternal influence on different aspects of dormancy behavior has been identified, its impact under field conditions and its relation to plant fitness has not been fully determined. This study examined maternal effects in Polygonum aviculare on release of seed primary dormancy, responses to alternating temperatures, induction into secondary dormancy, and field emergence patterns as influenced by changes in the sowing date and photoperiod experienced by the mother plant. Maternal effects were quantified using population threshold models that allowed us to simulate and interpret the experimental results. We found that regulation of dormancy in P. aviculare seeds by the maternal environment is instrumental for maximizing plant fitness in the field. This regulation operates by changing the dormancy level of seeds dispersed at different times (as a consequence of differences in the sowing dates of mother plants) in order to synchronize most emergence to the seasonal period that ultimately guarantees the highest reproductive output of the new generation. Our results also showed that maternal photoperiod, which represents a clear seasonal cue, is involved in this regulation.

Seed dormancy responses to temperature relate to Nothofagus species distribution and determine temporal patterns of germination across altitudes in Patagonia.

Seeds integrate environmental cues that modulate their dormancy and germination. Although many mechanisms have been identified in laboratory experiments, their contribution to germination dynamics in existing communities and their involvement in defining species habitats remain elusive. By coupling mathematical models with ecological data we investigated the contribution of seed temperature responses to the dynamics of germination of three Nothofagus species that are sharply distributed across different altitudes in the Patagonian Andes. Seed responsiveness to temperature of the three Nothofagus species was linked to the thermal characteristics of their preferred ecological niche. In their natural distribution range, there was overlap in the timing of germination of the species, which was restricted to mid-spring. By contrast, outside their species distribution range, germination was temporally uncoupled with altitude. This phenomenon was described mathematically by the interplay between interspecific differences in seed population thermal parameters and the range in soil thermic environments across different altitudes. The observed interspecific variations in seed responsiveness to temperature and its environmental regulation, constitute a major determinant of the dynamics of Nothofagus germination across elevations. This phenomenon likely contributes to the maintenance of patterns of species abundance across altitude by placing germinated seeds in a favorable environment for plant growth.

The role of seed water content for the perception of temperature signals that drive dormancy changes in Polygonum aviculare buried seeds.

Seedling emergence in the field is strongly related to the dynamics of dormancy release and induction of the seed bank, which is mainly regulated by soil temperature. However, there is limited information on how temperature-driven effects on dormancy changes are modulated by the seed hydration-level. We investigated the effect of seed water content (SWC) on the dormancy release and dormancy induction in Polygonum aviculare L. seeds. We characterised quantitatively the interaction between seed water content (SWC) and temperature through the measurement of changes in the lower limit temperature for seed germination (Tl) during dormancy changes for seeds with different SWC. These relationships were inserted in existing population-based threshold models and were run against field obtained data. The model considering SWC was able to predict P. aviculare field emergence patterns. However, failure to consider SWC led to overestimations in the emergence size and timing. Our results show that in humid temperate habitats, the occurrence of eventual water shortages during late-winter or spring (i.e. short periods of water content below 31% SWC) can affect soil temperature effects on seed dormancy, and might lead reductions in the emergence size rather than to significant temporal displacements in the emergence window. In conclusion, SWC plays an important role for the perception of temperature signals that drive dormancy changes in buried seeds.

Dormancy, a critical trait for weed success in crop production systems.

Agricultural practices exert selective forces on weed populations. As these practices change over time, weed adaptive traits also evolve, allowing weeds to persist in the new environment. However, only weeds having individuals showing the trait with adaptive significance will be able to cope with these changes, thus allowing a sub-population to be selected for persistence. In addition, changes in agricultural practices can select new weed species showing functional traits with characteristics adaptive to the modified system. Seed dormancy has long been recognized as a trait with enormous adaptive value to adjust weed biology to cropping systems. In this paper, we illustrate with examples of success and failure, the value of seed dormancy as a functional trait to cope with long-term changes in crop production systems. We show that successful outcomes are mostly related to the existence of sufficient variability for the functioning of physiological mechanisms that control dormancy characteristics as influenced by the agricultural environment. Presented examples illustrate how knowledge about the relationship that exists between agricultural practices and their selective pressure on seed dormancy can be instrumental in predicting changes in weed biotype dormancy characteristics or foreseeing the appearance of new weed species in future agricultural scenarios. © 2019 Society of Chemical Industry.

ABA inhibits germination but not dormancy release in mature imbibed seeds of Lolium rigidum Gaud.

Dormancy release in imbibed annual ryegrass (Lolium rigidum Gaud.) seeds is promoted in the dark but inhibited in the light. The role of abscisic acid (ABA) in inhibition of dormancy release was found to be negligible, compared with its subsequent effect on germination of dormant and non-dormant seeds. Inhibitors of ABA metabolism had the expected effects on seed germination but did not influence ABA concentration, suggesting that they act upon other (unknown) factors regulating dormancy. Although gibberellin (GA) synthesis was required for germination, the influence of exogenous GA on both germination and dormancy release was minor or non-existent. Embryo ABA concentration was the same following treatments to promote (dark stratification) and inhibit (light stratification) dormancy release; exogenous ABA had no effect on this process. However, the sensitivity of dark-stratified seeds to ABA supplied during germination was lower than that of light-stratified seeds. Therefore, although ABA definitely plays a role in the germination of annual ryegrass seeds, it is not the major factor mediating inhibition of dormancy release in imbibed seeds.

Involvement of ABA in induction of secondary dormancy in barley (Hordeum vulgare L.) seeds.

At harvest, barley seeds are dormant because their germination is difficult above 20 degrees C. Incubation of primary dormant seeds at 30 degrees C, a temperature at which they do not germinate, results in a loss of their ability to germinate at 20 degrees C. This phenomenon which corresponds to an induction of a secondary dormancy is already observed after a pre-treatment at 30 degrees C as short as 4-6 h, and is optimal after 24-48 h. It is associated with maintenance of a high level of embryo ABA content during seed incubation at 30 degrees C, and after seed transfer at 20 degrees C, while ABA content decreases rapidly in embryos of primary dormant seeds placed directly at 20 degrees C. Induction of secondary dormancy also results in an increase in embryo responsiveness to ABA at 20 degrees C. Application of ABA during seed treatment at 30 degrees C has no significant additive effect on the further germination at 20 degrees C. In contrast, incubation of primary dormant seeds at 20 degrees C for 48 and 72 h in the presence of ABA inhibits further germination on water similarly to 24-48 h incubation at 30 degrees C. However fluridone, an inhibitor of ABA synthesis, applied during incubation of the grains at 30 degrees C has only a slight effect on ABA content and secondary dormancy. Expression of genes involved in ABA metabolism (HvABA8'OH-1, HvNCED1 and HvNCED2) was studied in relation to the expression of primary and secondary dormancies. The results presented suggest a specific role for HvNCED1 and HvNCED2 in regulation of ABA synthesis in secondary seed dormancy.

Quantifying the sensitivity of barley seed germination to oxygen, abscisic acid, and gibberellin using a population-based threshold model.

Barley (Hordeum vulgare L.) seeds (grains) exhibit dormancy at maturity that is largely due to the presence of the glumellae (hulls) that reduce the availability of oxygen (O2) to the embryo. In addition, abscisic acid (ABA) and gibberellins (GAS) interact with O2 to regulate barley seed dormancy. A population-based threshold model was applied to quantify the sensitivities of seeds and excised embryos to O2, ABA, and GA, and to their interactive effects. The median O2 requirement for germination of dormant intact barley seeds was 400-fold greater than for excised embryos, indicating that the tissues enclosing the embryo markedly limit O2 penetration. However, embryo O2 thresholds decreased by another order of magnitude following after-ripening. Thus, increases in both permeability of the hull to O2 and embryo sensitivity to O2 contribute to the improvement in germination capacity during after-ripening. Both ABA and GA had relatively small effects on the sensitivity of germination to O2, but ABA and GA thresholds varied over several orders of magnitude in response to O2 availability, with sensitivity to ABA increasing and sensitivity to GA decreasing with hypoxia. Simple additive models of O2-ABA and O2-GA interactions required consideration of these O2 effects on hormone sensitivity to account for actual germination patterns. These quantitative and interactive relationships among O2, ABA, and GA sensitivities provide insight into how dormancy and germination are regulated by a combination of physical (O2 diffusion through the hull) and physiological (ABA and GA sensitivities) factors.

Pre-harvest Sprouting and Grain Dormancy in Sorghum bicolor: What Have We Learned?

The possibility of obtaining sorghum grains with quality to match the standards for a diversity of end-uses is frequently hampered by the susceptibility to pre-harvest sprouting (PHS) displayed by many elite genotypes. For these reasons, obtaining resistance to PHS is considered in sorghum breeding programs, particularly when the crop is expected to approach harvest maturity under rainy or damp conditions prevalence. As in other cereals, the primary cause for sprouting susceptibility is a low dormancy prior to crop harvest; in consequence, most research has focused in understanding the mechanisms through which the duration of dormancy is differentially controlled in genotypes with contrasting sprouting behavior. With this aim two tannin-less, red-grained inbred lines were used as a model system: IS9530 (sprouting resistant) and Redland B2 (sprouting susceptible). Redland B2 grains are able to germinate well before reaching physiological maturity (PM) while IS9530 ones can start to germinate at 40-45 days after pollination, well after PM. Results show that the anticipated dormancy loss displayed by Redland B2 grains is related reduced embryo sensitivity to abscisic acid (ABA) and increased levels of GA upon imbibition. In turn, transcriptional data showed that ABA signal transduction is impaired in Redland B2, which appears to have an impact on GA catabolism, thus affecting the overall GA/ABA balance that regulates germination. QTL analyses were conducted to test whether previous candidate genes were located in a dormancy QTL, but also to identify new genes involved in dormancy. These analyses yielded several dormancy QTL and one of them located in chromosome 9 (qGI-9) was consistently detected even across environments. Fine mapping is already in progress to narrow down the number of candidate genes in qGI-9.

On the hormonal nature of the stimulatory effect of high incubation temperatures on germination of dormant sorghum (S. bicolor) caryopses.

• High incubation temperatures (i.e. 30°C) stimulate the germination of dormant sorghum grains. To test the hormonal nature of this response, experiments were carried out with two varieties with contrasting dormancy at harvest: Redland B2 (low dormancy, high germination percentages attained under a wide incubation thermal range) and IS 9530 (high dormancy, high germination percentages attained only at 30°C). • Redland B2 grains with reduced GA content (paclobutrazol-treated) reached high germination temperatures (c. 100%) only when incubated at 30°C. By contrast, IS 9530 grains with reduced ABA content (fluridone-treated) reached 100% germination at 30, 25, 20 and 15°C. • Incubation temperatures did not alter embryo responsiveness to ABA, nor did it modify the pattern of changes in embryo ABA content throughout incubation. Low GA3 concentrations (0.1 µm) were required to totally overcome the inhibition imposed by ABA in embryos incubated at 30°C; by contrast, even the highest GA3 concentrations used (1000 µm) were not able to revert ABA inhibition in embryos incubated at 15°C. • These results show the hormonal nature of the stimulatory effect of high incubation temperatures, and suggest that this effect is mediated by an increase in tissue responsiveness to GAs.

Challenges facing seed banks and agriculture in relation to seed quality.

Seeds form a convenient vehicle for storage of germplasm, both for agricultural purposes and conservation of wild species. When required, seeds can be taken from storage and germinated, and plants can be propagated for the desired purpose, e.g., crop production or biome restoration. However, seed dormancy often interferes with stand establishment or industrial utilization in crops and germination of wild species. An anticipated termination of dormancy (i.e., before crop harvest) also occurs, with preharvest sprouting as a consequence. In order to overcome these problems, a better understanding of dormancy is required. This chapter is devoted to discuss the achievement of such understanding in problematic species.

Hypoxia interferes with ABA metabolism and increases ABA sensitivity in embryos of dormant barley grains.

Two mechanisms have been suggested as being responsible for dormancy in barley grain: (i) ABA in the embryo, and (ii) limitation of oxygen supply to the embryo by oxygen fixation as a result of the oxidation of phenolic compounds in the glumellae. The aim of the present work was to investigate whether hypoxia imposed by the glumellae interferes with ABA metabolism in the embryo, thus resulting in dormancy. In dormant and non-dormant grains incubated at 20 degrees C and in non-dormant grains incubated at 30 degrees C (i.e. when dormancy is not expressed), ABA content in the embryo decreased dramatically during the first 5 h of incubation before germination was detected. By contrast, germination of dormant grains was less than 2% within 48 h at 30 degrees C and embryo ABA content increased during the first hours of incubation and then remained 2-4 times higher than in embryos from grains in which dormancy was not expressed. Removal of the glumellae allowed germination of dormant grains at 30 degrees C and the embryos did not display the initial increase in ABA content. Incubation of de-hulled grains under 5% oxygen to mimic the effect of glumellae, restored the initial increase ABA in content and completely inhibited germination. Incubation of embryos isolated from dormant grains, in the presence of a wide range of ABA concentrations and under various oxygen tensions, revealed that hypoxia increased embryo sensitivity to ABA by 2-fold. This effect was more pronounced at 30 degrees C than at 20 degrees C. Furthermore, when embryos from dormant grains were incubated at 30 degrees C in the presence of 10 microM ABA, their endogenous ABA content remained constant after 48 h of incubation under air, while it increased dramatically in embryos incubated under hypoxia, indicating that the apparent increase in embryo ABA responsiveness induced by hypoxia was, in part, mediated by an inability of the embryo to inactivate ABA. Taken together these results suggest that hypoxia, either imposed artificially or by the glumellae, increases embryo sensitivity to ABA and interferes with ABA metabolism.

Expression analysis of a GA 20-oxidase in embryos from two sorghum lines with contrasting dormancy: possible participation of this gene in the hormonal control of germination.

The role of GAs in promoting seed germination is well known and experiments with seeds from different species have suggested the requirement of de novo synthesis of GAs upon imbibition for germination. There are also strong indications that the enhancement of GA synthesis is part of the mechanism through which environmental signals (i.e. light) induce germination. Since along the GA biosynthetic pathway, oxidation at C-20 carried out by GA 20-oxidases is thought to be a site of regulation, a cDNA clone encoding a GA 20-oxidase was isolated from embryos of sorghum (SbGA 20ox). Expression analysis of this gene in embryos within imbibed caryopses with low dormancy showed detectable amounts of the specific mRNA early upon incubation, increasing thereafter. In contrast, it remained barely detectable in embryos from dormant caryopses. Changes in endogenous GA4 levels were in agreement with those of SbGA 20ox mRNA, suggesting that GA production might be regulated differentially at the level of transcription of this gene. The expression of SbGA 20ox was enhanced in incubated embryos isolated from either type of caryopses, illustrating a physiological control exerted by the surrounding seed tissues on gene expression. The results also show that ABA leads to a suppression of transcription of this gene.