Seeds of the winter annual Bromus tectorum lose primary dormancy in summer and are poised to germinate rapidly in the autumn. If rainfall is inadequate, seeds remain ungerminated and may enter secondary dormancy under winter conditions. We quantified conditions under which seeds enter secondary dormancy in the laboratory and field and also examined whether contrasting B. tectorum genotypes responded differently to dormancy induction cues. The study also extends previous hydrothermal time models for primary dormancy loss and germination timing in B. tectorum by using similar models to account for induction and loss of secondary dormancy. Maximum secondary dormancy was achieved in the laboratory after 4 weeks at –1.0 MPa and 5°C. Seeds in the field became increasingly dormant through exposure to temperatures and water potentials in this range, confirming laboratory results. They were released from dormancy through secondary after-ripening the following summer. Different genotypes showed contrasting responses to dormancy induction cues in both laboratory and field. To examine secondary dormancy induction and release in the field in terms of hydrothermal time parameters, we first created a model that allowed mean base water potential (Ψb(50)) to vary while holding other hydrothermal time parameters constant, as in models for primary dormancy loss under dry conditions. The second model allowed all three model parameters to vary through time, to account for changes (e.g. hydrothermal time accumulation) that could occur simultaneously with dormancy induction in imbibed seeds. Shifts in Ψb(50) could explain most changes in dormancy status for seeds retrieved from the field, except during the short period prior to dormancy induction, when hydrothermal time was accumulating. This study illustrates that hydrothermal modelling, and specifically changes in Ψb(50), can be used to characterize secondary dormancy induction and loss in B. tectorum.

The transition from the dormant to the non-dormant state of a viable and mature seed can take place at low hydration by exposure to air-dry storage conditions (dry afterripening; AR). The events occurring during this loss of dormancy are of considerable physiological, ecological and agricultural interest. AR may be attributable to increased sensitivity to germination-stimulating factors and a widening of the temperature window for germination. Genetic, –omics and physiological studies on this mode of dormancy breaking provide support for a key role of the balance between gibberellin (GA) and abscisic acid (ABA) metabolism and sensitivity. Recent evidence also supports a possible role for ethylene (ET) in this complex signalling network that is necessary for AR implementation. However, hormone-independent signals, such as reactive oxygen species (ROS), nitrate () or nicotinamide adenine dinucleotide (NAD+), also appear to be involved. The way in which hormone- and non-hormone-signalling pathways affects each other (cross-talk) is still under study. This review provides updated information on the programmes that overcome seed dormancy. Thus, we have reviewed: (1) the –omic status in dry seeds; (2) the relationship between temperature and nitrate signalling and AR; (3) alterations in ABA/GA synthesis and signalling; (4) the action of hormone molecules other than ABA and GA (i.e. ET, salicylic and jasmonic acids); and (5) participation of reactive oxygen species (ROS), NAD+ and protein carbonylation. Taken together, the acquisition and implementation of dry AR involve a complex signalling network that is difficult to disentangle.

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