The role of light in regulating seed dormancy and germination

Seed dormancy is an adaptive trait in plants. Breaking seed dormancy determines the timing of germination and is, thereby essential for ensuring plant survival and agricultural production. Seed dormancy and the subsequent germination are controlled by both internal cues (mainly hormones) and environmental signals. In the past few years, the roles of plant hormones in regulating seed dormancy and germination have been uncovered. However, we are only beginning to understand how light signaling pathways modulate seed dormancy and interaction with endogenous hormones. In this review, we summarize current views of the molecular mechanisms by which light controls the induction, maintenance and release of seed dormancy, as well as seed germination, by regulating hormone metabolism and signaling pathways.     

Ecology of seed dormancy and germination in sedges (Carex)

The genus Carex, with its wide distribution and large number of species yet with a rather uniform life history, is a very convenient group for comparative studies of germination ecology at the generic level. The combination of a strict or conditional primary dormancy, a light requirement for germination, low germination at constant temperatures, a positive response to diurnal temperature fluctuations and an induction of secondary dormancy in late spring by increasing environmental temperatures are attributes that were found to be characteristics shared by almost all the Carex species investigated, though there was variation between species in the degree to which these characters were expressed. In almost all species, dormancy was broken by stratification at low temperatures, though few species gained the ability to germinate at temperatures <10 °C. There is evidence that long-term physiological changes and the structure of seed coats can play a decisive role in delaying germination. High dormancy levels were found mainly in Carices with large seeds (>0.9 mg), probably due to a thicker seed coat and hence a higher resistance to germination. Differences in primary dormancy between sedges of various habitats could not be established. However, there was a tendency for temperature limits to be low in forest sedges. Many species of wetlands and open sites showed a greater capability to respond to fluctuating temperatures than species of dry sites. These dormancy and germination traits not only enable the accumulation of seeds in the soil, but also constitute seasonal seed regeneration strategies that rely on the high longevity of seeds and the formation of persistent seed banks. Temperate Carices are mainly adapted to exploit the temporally and spatially infrequent occurrence of canopy gaps that become available only in late spring or early summer, whereas the colonization of gaps at the beginning of the vegetation period is largely prevented by a high temperature requirement for germination. Many of the dormancy and germination characteristics of Carices are important in Cyperaceae generally. A greater diversity of germination responses, however, can be found in the related families, Juncaceae and Poaceae. Our present knowledge is not sufficient to determine unequivocally whether a phylogenetic component contributes significantly to the germination behaviour of the genus Carex, but certain tendencies are clearly indicated.     

两种结缕草种子休眠及萌发特性

通过测定兰引Ⅲ号结缕草和青岛结缕草休眠种子和解除休眠种子的吸水率、呼吸强度和脱氢酶活性等萌发生理指标,探讨了结缕草种子的休眠类型。试验结果表明,解除休眠的结缕草种子的吸水率大于未处理的种子。解除休眠种子的呼吸强度、脱氢酶活性都有较大幅度的增长,显著高于未处理。结缕草种子萌发期需30～35℃高温和充足光照。两种结缕草种子的颖壳、种皮的透性障碍及种子内存在发芽抑制物质是导致种子休眠的主要原因,属于混合休眠类型。     

Variation in seed dormancy and germination among populations of Silybum marianum (Asteraceae)

Milk thistle (Silybum marianum) is a medicinal plant; however, lack of consistency in past dormancy studies has hindered propagation of this species from seeds. We tested the germination responses of freshly harvested and after-ripened (stored for 2 and 7 months; 25°C at 50% relative humidity) seeds from three populations (P1, P2 and P3) in Iran at varying constant or alternating temperatures, with or without GA₃ and in light and continuous darkness. No germination occurred in freshly harvested seeds incubated at any condition without GA₃ application, indicating that all the seeds were dormant. Seeds from P1 and P2, which developed under relatively dry, warm conditions, germinated over a wider range of temperatures after 2 months of dry storage, indicating type 6 of non-deep physiological dormancy (PD). Seeds from P3, which developed under relatively wet, cool conditions, incubated at constant temperatures (especially on GA₃), exhibited an increase in maximum temperature for germination, indicating type 1 of non-deep PD. Light improved germination of after-ripened seeds, and GA₃ application substituted for the light requirement for germination. This is the first report that environmental conditions during seed development may be correlated with differences in the type of non-deep PD. We conclude that milk thistle seeds are positively photoblastic and photodormant and the germination responses of after-ripened seeds from different populations are different under darkness. Therefore, the impacts of genetic differences and maternal effects on the induction of dormancy during seed development should be considered in attempts to domesticate this medicinal plant.     

Effects of seed size and frugivory degree on dispersal by Neotropical frugivores

Large vertebrates are important elements of mutualistic interactions and provide positive impacts on plant population and community dynamics. Despite the increasing interest on vertebrate frugivory we are still not able to disentangle the real contribution of seed dispersal to Neotropical forest functioning. Consuming fruits does not imply effective seed dispersal and many variables, such as seed size and animal diet, may influence the outcome of plant-animal interactions. Here, we performed a comprehensive literature search on seed dispersal by Neotropical vertebrates (with a focus on primates) to closely approach their role as seed dispersers, hypothesizing frugivory degree and seed size as main drivers of fruit handling behavior and diversity of dispersed seeds. We found that the great majority of seeds manipulated by Neotropical primates, with exception to the seed predators pitheciins, were swallowed and passed intact through their gut. Larger seeds (>12 mm) tended of being ingested exclusively by primates and other large vertebrates, such as tapirs and peccaries. Furthermore, primate feeding guild had a great influence on the richness and sizes of seeds dispersed, as primarily frugivores dispersed more species and had higher probabilities of ingesting larger seeds when compared to other feeding guilds. Organizing available knowledge and filling the main knowledge gaps allowed us to validate common sense assumptions and ultimately draw new conclusions about the role played by primates together with other major frugivores in Neotropical forests.     

The control of seed germination in Trollius ledebouri A model of seed dormancy

Treatment of Trollius ledebouri seeds with gibberellins A₄+A₇ promotes germination. The efficacy of the treatment is dependent upon the duration of imbibition in distilled water prior to GA₄₊₇ application. Presoaking increases both the final percentage germination attained and also its rate of achievement. No presoaking effect is exhibited by seeds induced to germinate by testa removal in the absence of GA₄₊₇. Active washing of Trollius seeds enhances the presoaking effect and the eluent from washed seeds is inhibitory to germination. The results support the hypothesis that the presoaking effect exhibited by Trollius is the result of the leaching of a germination inhibitor from the seeds which is antagonistic to GA₄₊₇. Additionally, treatment of Trollius seeds with the gibberellin-biosynthesis inhibitor (2-chloroethyl)-trimethylammonium chloride (CCC) prior to testa removal retards germination. The inhibitory effect of CCC on germination is overcome by GA₄₊₇. Although CCC inhibits embryo growth during the presoaking of intact seeds, it does not affect the increased sensitivity of presoaked seeds to GA₄₊₇. Therefore, although endogenous gibberellins may be involved in the germination process, they do not contribute to the presoaking phenomenon. The expansion of isolated endosperm tissue is not affected by CCC. However, the chemical markedly inhibits endosperm expansion in intact seeds and implicates the embryo as both the site of production of the germination inhibitor and of gibberellin. These results are discussed in relation to previous studies and a model is presented to account for the characteristics of germination in Trollius.Abbreviations GA gibberellin - CCC (2-chloroethyl)-trimethylammonium chloride     

生长素调控种子的休眠与萌发

植物种子的休眠与萌发,是植物生长发育过程中的关键阶段,也是生命科学领域的研究热点。种子从休眠向萌发的转换是极为复杂的生物学过程,由外界环境因子、体内激素含量及信号传导和若干关键基因协同调控。大量研究表明,植物激素脱落酸(Abscisic acid, ABA)和赤霉素(Gibberellin acid, GA)是调控种子休眠水平,决定种子从休眠转向萌发的主要内源因子。ABA与GA在含量和信号传导两个层次上的精确平衡,确保了植物种子能以休眠状态在逆境中存活,并在适宜的时间启动萌发程序。生长素(Auxin)是经典植物激素之一,其对向性生长和组织分化等生物学过程的调控已有大量研究。但最近有研究证实,生长素对种子休眠有正向调控作用,这表明生长素是继ABA之后的第二个促进种子休眠的植物激素。本文在回顾生长素的发现历程、阐释生长素体内合成途径及信号传导通路的基础上,重点综述了生长素通过与ABA的协同作用调控种子休眠的分子机制,并对未来的研究热点进行了讨论和展望。     

Temperature effects on dormancy levels and germination in temperate forest sedges (Carex)

The effects of stratification temperatures and burial in soil on dormancy levels of Carex pendula L. and C. remota L., two spring-germinating perennials occurring in moist forests, were investigated. Seeds buried for 34 months outdoors, and seeds stratified in the laboratory at temperatures between 3 and 18 °C for periods between 2 and 28 weeks, were tested over a range of temperatures. Seeds of the two species responded similarly to stratification treatments, except for an absolute light requirement in C. pendula. Primary dormancy was alleviated at all stratification temperatures, but low temperatures were more effective than higher ones . (≥ 12 °C). Dormancy induction in non-dormant seeds kept at 5 °C occurred when seeds were subsequently exposed to 18 °C. Dormancy was not induced by a transfer to lower temperatures. Buried seeds of both species exhibited seasonal dormancy cycles with high germination from autumn to spring and low germination during summer. Temperatures at which the processes of dormancy relief and of dormancy induction occurred, overlapped to a high degree. Whether, and when, dormancy changes occurred depended on test conditions. The lower temperature limit for germination (> 10%) was 9 °C in C. remota and 15 °C in C. pendula. Germination ceased abruptly above 36 °C. Germination requirements and dormancy patterns suggest regeneration from seed in late spring and summer at disturbed, open sites (forest gaps) and the capability to form long, persistent seed banks in both species.     

Temperature controls seed germination and dormancy in the European woodland herbaceous perennial Erythronium dens-canis (Liliaceae)

We examined the germination ecology and the temperature requirements for germination of Erythronium dens-canis, under both outdoor and laboratory conditions. E. dens-canis is a spring flowering woodland geophyte widely distributed across Europe. Germination phenology, including embryo development and radicle and cotyledon emergence, were investigated in a natural population growing in Northern Italy. Immediately after harvest, seeds of E. dens-canis were either sown on agar in the laboratory under simulated seasonal temperatures or placed in nylon mesh sachets and buried in the wild. Embryos, undifferentiated at the time of seed dispersal, grew during summer and autumn conditions in the laboratory and in the wild, culminating in radicle emergence in winter when temperatures fell to ≈ 5 °C. Emergence of cotyledons did not occur immediately after radicle emergence, but was delayed until the end of winter. Laboratory experiments showed that temperature is the main factor controlling dormancy and germination, with seeds becoming non-dormant only when given warmth, followed by cold stratification. Unlike seeds of E. dens-canis that germinate in winter, in other Erythronium species radicle emergence occurs in autumn, while in some it is delayed until seeds are transferred from winter to spring conditions. Our results suggest that there is genetic and environmental control of the expression of seed dormancy amongst Erythronium species, which is related to local climate.     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

萌发前处理和种子大小对瓜拉那(Paullinia cupana var.sorbilis)种子萌发的影响

为了增加对瓜拉那(Paullinia cupana var.sorbilis)种子萌发特性的了解,寻求提高瓜拉那种子萌发的最佳条件,本研究用控制实验,研究了瓜拉那种子去/不去假种皮、不同种子大小和酸处理下的萌发行为.结果表明,去假种皮且酸处理的大种子发芽率最高,为100%;不去假种皮且无酸处理的小种子基本不萌发;去假种皮种子发芽率显著高于未去假种皮种子;种子大小对萌发率有显著影响,大种子比小种子趋于有更高的萌发率;酸处理能进一步提高去假种皮种子萌发率.     

Morphological side effects of using gibberellic acid to induce germination: consequences for the study of seed dormancy

To assess the evolutionary significance of persistent seed banks, phenotypes of naturally germinating seeds must be compared with those that remain dormant under the same environmental conditions. Dormant seeds can often be induced to germinate by application of gibberellic acid (GA). However, this method is valid only if there are no phenotypic “side effects” of GA that could confound comparisons between dormant and naturally germinating seeds. We examined this assumption in Lesquerella fendleri, a short-lived perennial mustard of the desert Southwest. We exposed 3840 seeds from 16 maternal sibships to two different GA treatments (0 or 1 g/L GA) in two different germination environments (greenhouse and growth chamber), and measured germination and postgermination traits. As expected, application of GA increased germination. GA also had strong and long-lasting effects on seedling morphology. Seeds that received GA developed into seedlings that were taller, with fewer but longer leaves, than seeds that did not receive GA. Effects of GA on both dormancy and postgermination traits varied among maternal sibships. Our results indicate that for this species and this concentration of GA, morphological effects can be substantial. Further study is required to determine whether such side effects are found for lower concentrations of GA, or under conditions that encourage faster seedling growth. Nonetheless, the present results illustrate the importance of testing potential confounding effects of GA in studies of the evolution of seed dormancy and its influence on postgermination traits.     

Seed dormancy and germination (tertiary level biology)

Book review

Seed dormancy and germination (tertiary level biology)J W Bradbeer, Glasgow: Blackie &; Son 1988. x + 146 pages. £23.00 hardback; £10.95 paperback. ISBN 0-216 91635-6; ISBN 0-216-91636-4 PbK     

Influence of the testa on seed dormancy, germination, and longevity in Arabidopsis

The testa of higher plant seeds protects the embryo against adverse environmental conditions. Its role is assumed mainly by controlling germination through dormancy imposition and by limiting the detrimental activity of physical and biological agents during seed storage. To analyze the function of the testa in the model plant Arabidopsis, we compared mutants affected in testa pigmentation and/or structure for dormancy, germination, and storability. The seeds of most mutants exhibited reduced dormancy. Moreover, unlike wild-type testas, mutant testas were permeable to tetrazolium salts. These altered dormancy and tetrazolium uptake properties were related to defects in the pigmentation of the endothelium and its neighboring crushed parenchymatic layers, as determined by vanillin staining and microscopic observations. Structural aberrations such as missing layers or a modified epidermal layer in specific mutants also affected dormancy levels and permeability to tetrazolium. Both structural and pigmentation mutants deteriorated faster than the wild types during natural aging at room temperature, with structural mutants being the most strongly affected.     

Fungal spores: dormancy, germination, chemical composition, and role in biotechnology (review)

This review is focused on one of the stages of ontogenesis distinctive by its particular tolerance to the action of unfavorable factors and ability to retain the genomic material for a long period of time, i.e., fungal spores. The major part is devoted to the characterization of the specific stage typical for spores, which is called dormancy. Data are presented characterizing the carbohydrate and lipid composition of spores, with special attention being paid to the role of carbohydrate protectors, in particular, trehalose and mannite, as well as to the role of rafts in the process of sporogenesis. The role of special compounds called autoinhibitors and autostimulators in the process of exit from dormancy is discussed. The final section deals with the role of spore seeding material in biotechnological processes. Data on the correlation between the chemical composition of spores, their ability to remain dormant, and the germination process are considered. Special biotechnological approaches are presented for the first; they allow for the preservation of the germinating ability of spores, intensification of sporogenesis, changes in the ratio of final fermentation products, and an increase in their yield.     

MicroRNA在种子发育、休眠与萌发过程中的作用

植物microRNA(miRNA)是一类小分子非编码RNA,对植物的生长发育发挥着重要调控作用。种子发育、休眠与萌发是植物生命进程中的重要阶段。在这一阶段内,种子受各种环境因子及内源激素调控,并且不同植物种子具有不同发育及休眠特性。随着人们对种子发育、休眠及萌发机理的探究,越来越多miRNA被鉴定,它们能够基于植物激素信号传导、抗氧化作用、关键转录因子调控等途径参与种子形态建成、物质代谢及各种胁迫响应。本文主要综述了近年来植物miRNA的形成及调控机理,以及在种子发育、休眠及萌发过程中发挥的调控作用,旨在为今后的研究方向提供参考。     

Discovering the type of seed dormancy and temperature requirements for seed germination of Gentiana lutea L. subsp. lutea (Gentianaceae)

Aims There are a number of mechanisms that regulate germination; among these, seed dormancy, one of the most important, is an adaptative mechanism in plants to promote survival by dispersing germination in space and time until environmental conditions are favourable for germination. The main goals of this study were to determine the temperature requirements for seed dormancy release and germination of Gentiana lutea subsp. lutea, to identify the class and level of seed dormancy and to suggest an optimal germination protocol.Methods Seeds belonging to two different localities were subjected to various pre-treatments, including cold stratification (0 and 5°C), warm stratification (25/10°C) and different combinations of these, and then incubated at a range of constant temperatures (5–25°C) and 25/10°C. Embryo growth during pre-treatments and incubation conditions were assessed at different times by measuring the embryo to seed length ratio (E:S ratio). The final germination percentage (FGP) and the germination rate (t 50) were calculated.Important findings Fleshy mature seeds of G. lutea subsp. lutea have linear underdeveloped embryos. Cold stratification at 0°C was effective in overcoming the physiological dormancy (PD) and promoted embryo growth and subsequent germination. After cold stratification at 0°C, both the root and the shoot emerged readily under a wide range of temperatures. G. lutea subsp. lutea seeds showed an intermediate complex morphophysiological dormancy (MPD). As regards the optimal germination protocol for this taxon, we suggest a period of cold stratification at ca. 0°C followed by seed incubation at 10–20°C. The optimal germination temperatures found for seeds of this taxon, as well as its pre-chilling requirement at 0°C, suggest that it is well adapted to a temperate climate; this behavior highlights an increasing threat from global warming for G. lutea, which could reduce the level of natural emergence in the field, prejudicing also the long-term persistence of the natural populations in Sardinia.