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Species with physically dormant (PY) seeds make up over 25% of plant species in a number of ecologically important ecosystems around the globe, such as savannah and Mediterranean shrublands. Many of these ecosystems are subject to temporally stochastic events, such as fire and drought; but are in areas projected to experience some of the most extreme climatic changes in the future. Given the importance of PY in controlling germination timing for successful recruitment, we ask how plastic the PY trait is, and if changes to the maternal environment from climate change could alter recruitment. This review focuses on: (1) the evidence for inter- and intraspecific variation in PY; (2) the genetic, maternal and environmental controls involved; and (3) the ecological consequences of (1) and (2) above. Evidence for (within-community) interspecific variation in conditions required to break PY is strong, but for intraspecific variation evidence is contradictory and limited by a paucity of studies. Identifying controllers of variation in PY is complex, there is some suggestion that conditions of the maternal environment may be important, but no consensus on the nature of effects. The implications of PY plasticity for the persistence of seed banks, species and communities under climate change are discussed. We highlight a number of key knowledge gaps, such as a lack of research estimating the components of variation in non-agricultural species, and identify a suite of seed attributes relevant to understanding the potential impacts of climate change on the population dynamics of PY species in the future.
Physiological dormancy has been described as a physiological inhibiting mechanism that prevents radicle emergence. It can be caused by the embryo (embryo dormancy) as well as by the structures that cover the embryo. One of its functions is to time plant growth and reproduction to the most optimal season and therefore, in nature, dormancy is an important adaptive trait that is under selective pressure. Dormancy is a complex trait that is affected by many loci, as well as by an intricate web of plant hormone interactions. Moreover, it is strongly affected by a multitude of environmental factors. Its induction, maintenance, cycling and loss come down to the central paradigm, which is the balance between two key hormonal regulators, i.e. the plant hormone abscisic acid (ABA), which is required for dormancy induction, and gibberellins (GA), which are required for germination. In this review we will summarize recent developments in dormancy research (mainly) in the model plant Arabidopsis thaliana, focusing on two key players for dormancy induction, i.e. the plant hormone ABA and the DELAY OF GERMINATION 1 (DOG1) gene. We will address the role of ABA and DOG1 in relation to various aspects of seed dormancy, i.e. induction during seed maturation, loss during dry seed afterripening, the rehydrated state (including dormancy cycling) and the switch to germination.
As in other cultivated species, dormancy can be seen as a problem in cereal production, either due to its short duration or to its long persistence. Indeed, cereal crops lacking enough dormancy at harvest can be exposed to pre-harvest sprouting damage, while a long-lasting dormancy can interfere with processes that rely on rapid germination, such as malting or the emergence of a uniform crop. Because the ancestors of cereal species evolved under very diverse environments worldwide, different mechanisms have arisen as a way of sensing an appropriate germination environment (a crucial factor for winter or summer annuals such as cereals). In addition, different species (and even different varieties within the same species) display diverse grain morphology, allowing some structures to impose dormancy in some cereals but not in others. As in seeds from many other species, the antagonism between the plant hormones abscisic acid and gibberellins is instrumental in cereal grains for the inception, expression, release and re-induction of dormancy. However, the way in which this antagonism operates is different for the various species and involves different molecular steps as regulatory sites. Environmental signals (i.e. temperature, light quality and quantity, oxygen levels) can modulate this hormonal control of dormancy differently, depending on the species. The practical implications of knowledge accumulated in this field are discussed.
Many studies have claimed that fire acts as the chief ecological factor cueing dormancy break in seeds with a water-impermeable seed coat, i.e. physical dormancy (PY), in Mediterranean ecosystems. However, a proposal is made that seasonal temperature changes must be viewed as more meaningful dormancy-breaking cues because: (1) fire is erratic and may break PY in seasons during which seedlings cannot complete their life cycle; (2) fire may not occur for long periods, thereby only providing an opportunity for dormancy break and germination once in every several years; and (3) if fire opens the specialized anatomical structures called ‘water gaps’, in seconds, their evolutionary role of detecting environmental conditions becomes irrational. Although fire breaks dormancy in a proportion of seeds, given the risk of seed mortality and the post-fire environment providing cues for dormancy break, it is suggested that fire might possibly be an exaptation.
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.
Seeds of Sisymbrium officinale display physiological dormancy and require nitrate to germinate. Rupture of the testa precedes radicle protrusion through the endosperm (germination sensu stricto). While both endosperm rupture and testa rupture (TR) required nitrate, endosperm rupture was fully inhibited by abscisic acid (ABA) but TR was not inhibited. The gibberellic acid (GA)-synthesis inhibitor paclobutrazol prevented TR, which was reverted by exogenous GA4 but not by nitrate. The orientation of TR was transverse, which prompted the question whether seeds elongate prior to radicle protrusion, concurrent with an increase in water content. Between 9 h and 1 d no increase in length or water content was observed. During incubation in ABA the length of imbibed seeds without TR did not increase between 1 and 5 d, whereas nitrate added to ABA induced TR and a 94% increase in length. At the same time the water content of seeds without TR increased by 18%, while the water content of seeds with TR increased by 38%. Length and water content were correlated in a single-seed analysis for seeds with TR, but not for seeds without TR. Increased length was also observed in Arabidopsis seeds with nitrate-induced TR. These results indicate that prior to endosperm rupture dormancy release by nitrate is accompanied by TR, seed elongation and an increase in water content. A new multiphasic model is proposed for the imbibition curve, where the second phase of the classical triphasic curve is split into three sub-phases, of which phases IIB and IIC are associated with TR.
Temperature is a key factor affecting both dormancy and germination. In non-dormant seeds, when temperature is within the thermal range permissive for germination, it just regulates germination velocity, while in seeds presenting dormancy it can also be affecting dormancy level, dormancy termination and the expression of dormancy itself. This dual effect of temperature on dormancy and germination often leads to misinterpretation of obtained germination results and confounds the analysis of temperature effects in seed populations presenting some degree of dormancy. In the present paper we discuss the effect of temperature in the regulation of dormancy level and its implications in dormancy expression, as an attempt to construct a conceptual framework that allows distinguishing between the effects of temperature on dormancy and germination. Finally, we present examples of how a better understanding of these effects could help us to interpret the mixed effects of temperature on both processes during incubation of seeds presenting dormancy.
Seeds have evolved to be highly efficient environmental sensors that respond not only to their prevailing environment, but also their environmental history, to regulate dormancy and the initiation of germination. In the present work we investigate the combined impact of a number of environmental signals (temperature, nitrate, light) during seed development on the mother plant, during post-shedding imbibition and during prolonged post-shedding exposure in both dry and imbibed states, simulating time in the soil seed bank. The differing response to these environments was observed in contrasting winter (Cvi, Ler) and summer (Bur) annual Arabidopsis ecotypes. Results presented show that environmental signals both pre- and post-shedding determine the depth of physiological dormancy and therefore the germination response to the ambient environment. The ecotype differences in seed response to ambient germination conditions are greatly enhanced by seed maturation in different environments. Further variation in response develops following shedding when seeds do not receive the full complement of environmental signals required for germination and enter the soil seed bank in either dry or imbibed states. Species seed dormancy characteristics cannot therefore be easily defined, as seed dormancy is a dynamic state subject to within-species adaptation to local environments.
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.
Knowledge about the hormonal control of grain dormancy and dormancy loss is essential in wheat, because low grain dormancy at maturity is associated with the problem of pre-harvest sprouting (PHS) when cool and rainy conditions occur before harvest. Low GA (gibberellin A) hormone sensitivity and high ABA (abscisic acid) sensitivity were associated with higher wheat grain dormancy and PHS tolerance. Grains of two PHS-tolerant cultivars were very dormant at maturity, and insensitive to GA stimulation of germination. More PHS-susceptible cultivars were less sensitive to ABA inhibition of germination, and were either more GA sensitive or germinated efficiently without GA at maturity. As grain dormancy was lost through dry afterripening or cold imbibition, grains first gained GA sensitivity and then lost ABA sensitivity. These changes in GA and ABA sensitivity can serve as landmarks defining stages of dormancy loss that cannot be discerned without hormone treatment. These dormancy stages can be used to compare different cultivars, seed lots and studies. Previous work showed that wheat afterripening is associated with decreasing ABA levels in imbibing seeds. Wheat grain dormancy loss through cold imbibition also led to decreased endogenous ABA levels, suggesting that reduced ABA signalling is a general mechanism triggering dormancy loss.
Determining the phylogenetic and biogeographic distribution of physical dormancy remains a major challenge in germination ecology. Here, our goal was to describe a novel water-impermeable seed coat mechanism causing physical dormancy (PY) in the seeds of Chaetostoma armatum (Melastomataceae). Although seed coat permeability tests indicated a significant increase in seed weight after soaking in distilled water, anatomical and dye-tracking analyses showed that both water and dyes penetrated the seed coat but not the embryo, which remained in a dry state. The water and dye penetrated the lumen of the exotestal cells, which have a thin outer periclinal face and thickened secondary walls with U-shaped phenolic compounds. Because of this structure, water and dye do not penetrate the inner periclinal face of the exotestal cells, indicating PY. Puncturing the seeds increased germination more than tenfold compared to that of the control, but GA3 did not increase germination further. A significant fraction of the seeds did not germinate after puncturing, indicating that embryos are also physiologically dormant (PD). This paper constitutes the first report of the water-impermeable seed coat in the Myrtales and the first report of physiophysical (PD+PY) dormancy in a shrub from a tropical montane area.
The Araliaceae is known to have seeds with underdeveloped embryos that must grow prior to radicle emergence, and thus they have morphological (MD) or morphophysiological (MPD) dormancy. Araliaceae is one of about 15 families with woody species in the tropical montane zone, and in Hawaii 15 species occur in the montane. Our purpose was to determine if seeds of the Hawaiian Araliaceae species Cheirodendron trigynum subsp. trigynum have MD or MPD and, if MPD, what level. In a move-along experiment, some seeds were incubated continuously at 15/6, 20/10 or 25/15°C, while others were moved sequentially from low to high or from high to low temperature regimes. Germination percentages and embryo growth were monitored. Also, the effects of cold and warm stratification on dormancy break were determined. Seeds had physiological dormancy (PD) in addition to small embryos that grew prior to germination, and thus MPD. PD was broken slowly ( ≥ 12 weeks), after which embryos grew rapidly, followed by root and shoot emergence. Embryos grew at temperatures suitable for warm stratification; thus, seeds have Type 1 non-deep simple MPD; the dormancy formula is C1bBb. Seeds from Oahu germinated to 94–100% at 15/6, 20/10 and 25/15°C, while those from the Big Island germinated to high percentages only at 15/6 and 20/10°C. Temperature shifts improved germination of seeds from the Big Island, and movement from either low to high or from high to low temperature regimes was effective in promoting germination. This is the first report of non-deep simple MPD in the Araliaceae.
Convallariamajalis has double dormancy and hypogeal germination, but no information is available on embryo growth or on the effects of light and gibberellic acid (GA3) on germination in this genus. Therefore, we investigated embryo growth and other germination features in seeds of C. keiskei and compared the data with those of Trillium camschatcense in another study. Until now, in seeds with double dormancy, embryo growth and germination (epigeal) have been studied in detail only for seeds of T. camschatcense. Phenology of embryo growth and emergence of cotyledonary petiole/root (hereafter root) and shoot in seeds of C. keiskei were monitored outdoors. Effects of temperature, light and GA3 on embryo growth and root and shoot emergence were tested under laboratory conditions. Roots emerged the first spring following seed dispersal in autumn. The embryo grew soon after root emergence, and germination was hypogeal. Seeds with an emerged root formed buds from which a shoot (leaf) emerged above ground during the second spring. Alternating temperatures and light had negative effects on root emergence, and GA3 did not substitute for cold stratification in root emergence. Seeds of C. keiskei have double dormancy, but it differs from that in T. camschatcense. Based on differences in embryo growth before (T. camschatcense) versus after (C. keiskei) root emergence, and on epigeal (T. camschatcense) versus hypogeal (C. keiskei) germination, we suggest that two types of deep simple double morphophysiological dormancy (MPD) be recognized. Since embryo growth in C. keiskei does not fit the standard definition of MPD, we propose to expand this definition.
Dormancy has evolved in plants to restrict germination to favourable growth seasons. Seeds from most crop plants have low dormancy levels due to selection for immediate germination during domestication. Seed dormancy is usually not completely lost and low levels are required to maintain sufficient seed quality. Brassica napus cultivars show low levels of primary seed dormancy. However, B. napus seeds are prone to the induction of secondary dormancy, which can lead to the occurrence of volunteers in the field in subsequent years after cultivation. The DELAY OF GERMINATION 1 (DOG1) gene has been identified as a major dormancy gene in the model plant Arabidopsis thaliana. DOG1 is a conserved gene and has been shown to be required for seed dormancy in various monocot and dicot plant species. We have identified three B. napus genes with high homology to AtDOG1, which we named BnaA.DOG1.a, BnaC.DOG1.a and BnaC.DOG1.b. The transcripts of these genes could only be detected in seeds and showed a similar expression pattern during seed maturation as AtDOG1. In addition, the BnaDOG1 genes showed enhanced transcript levels after the induction of secondary dormancy. These results suggest a role for DOG1 in the induction of secondary dormancy in B. napus.
Seed dormancy can prevent germination under unfavourable conditions that reduce the chances of seedling survival. Freshly harvested seeds often have strong primary dormancy that depends on the temperature experienced by the maternal plant and which is gradually released through afterripening. However, seeds can be induced into secondary dormancy if they experience conditions or cues of future unfavourable conditions. Whether this secondary dormancy induction is influenced by seed-maturation conditions and primary dormancy has not been explored in depth. In this study, we examined secondary dormancy induction in seeds of Arabidopsis thaliana matured under different temperatures and with different levels of afterripening. We found that low water potential and a range of temperatures, from 8°C to 35°C, induced secondary dormancy. Secondary dormancy induction was affected by the state of primary dormancy of the seeds. Specifically, afterripening had a non-monotonic effect on the ability to be induced into secondary dormancy by stratification; first increasing in sensitivity as afterripening proceeded, then declining in sensitivity after 5 months of afterripening, finally increasing again by 18 months of afterripening. Seed-maturation temperature sometimes had effects that were independent of expressed primary dormancy, such that seeds that had matured at low temperature, but which had comparable germination proportions as seeds matured at warmer temperatures, were more easily induced into secondary dormancy. Because seed-maturation temperature is a cue of when seeds were matured and dispersed, these results suggest that the interaction of seed-maturation temperature, afterripening and post-dispersal conditions all combine to regulate the time of year of seed germination.