Seed dormancy and the control of germination

Seed dormancy is an innate seed property that defines the environmental conditions in which the seed is able to germinate. It is determined by genetics with a substantial environmental influence which is mediated, at least in part, by the plant hormones abscisic acid and gibberellins. Not only is the dormancy status influenced by the seed maturation environment, it is also continuously changing with time following shedding in a manner determined by the ambient environment. As dormancy is present throughout the higher plants in all major climatic regions, adaptation has resulted in divergent responses to the environment. Through this adaptation, germination is timed to avoid unfavourable weather for subsequent plant establishment and reproductive growth. In this review, we present an integrated view of the evolution, molecular genetics, physiology, biochemistry, ecology and modelling of seed dormancy mechanisms and their control of germination. We argue that adaptation has taken place on a theme rather than via fundamentally different paths and identify similarities underlying the extensive diversity in the dormancy response to the environment that controls germination.     

Seed dormancy and germination in Dodonaea viscosa (Sapindaceae) from south-western Saudi Arabia

Dodonaea viscosa (Sapindaceae) is widespread in the mountainous highlands of the southwestern part of Kingdom of Saudi Arabia, where it is a medicinally important species for the people in Saudi Arabia. Seeds of this species were collected from Mount Atharb in Al-Baha region, at an altitude of 2100 m. The aims of this study were to determine if the seeds of D. viscosa have physical dormancy (i.e. a water-impermeable seed coat) and, if so, what treatments would break dormancy, and what conditions promote germination after dormancy has been broken. The dormancy-breaking treatments included: soaking of seeds in concentrated sulfuric acid (H₂SO₄) for 10 min, immersion in boiling water for 10 min and exposure to 50 °C for 1 min. After seeds had been pre-treated with H₂SO₄, to break dormancy, they were incubated at constant temperatures from 5 to 35 °C, under 12-h photoperiods or in continuous darkness, and germination recorded. Salinity tolerance was investigated by incubating acid-scarified seeds in different concentrations of mM NaCl in the light at 25 °C.Untreated seeds had low final germination 30%. Seeds that had been acid-scarified, immersed in boiling water or exposed to 50 °C all achieved 91% subsequently when incubated at 25 °C. Thus, seeds of this species in Saudi Arabia have physical dormancy, which can be broken by all three treatments designed to increase the permeability of the testa. After pre-treatment, there was a broad optimum constant temperature for germination that ranged between 5 and 25 °C but germination was inhibited by higher temperatures (30 and 35 °C). Light had little effect on this germination response. Scarified seeds were also sensitive to salinity, with the highest germination in distilled water and complete inhibition in 400 mM NaCl. Seeds that failed to germinate in saline treatments were mostly able to germinate on transfer to distilled water, suggesting osmotic inhibition.     

Seed germination and dormancy responses to acetone condensation products

Acetone condensation products reported to have growth regulatingproperties were tested for their effects on seed germination.The active compounds—isoxylitones, isophorone, diacetonealcohol and phorone inhibited seed germination. The degree ofinhibition depends on the particular compound, its concentration,and the light and temperature environment of the imbibed seeds.Germination studies involving light quality, light energiesand temperature, and light microscope studies indicated little,if any, penetration of the chemicals into the seed but rathersurface effects. The results suggest that the chemicals inducedmembrane impermeability which was partially relieved by redlight and by a brief high-temperature treatment. (Received May 10, 1976; )     

旱生灌木岷谷木蓝种子的休眠与萌发特征

岷谷木蓝(Indigofera lenticellata)是横断山区干旱河谷广泛分布的豆科灌木.本文对其种子的休眠与萌发特征进行了研究.结果表明:岷谷木蓝种子具有物理休眠,其萌发困难的原因在于种皮的不透水性;种子的硬实比率高(85%),针刺种皮和沙摩处理能有效打破种子休眠,提高发芽率,其中沙摩7 h最为快速有效.岷谷木蓝种子发芽温度范围较广,10℃～30℃均可发芽,最适温度范围为20℃～30℃,低温不利于种子萌发.岷谷木蓝具有一定的抗旱性,在聚乙二醇6000(PEG)溶液模拟水分胁迫条件下,种子在25% PEG胁迫下仍能达到100%的发芽率,但水分胁迫对种子发芽有一定的延缓作用,发芽率和发芽速度降低,发芽周期和发芽准备期增长,发芽高峰期滞后.其中对于发芽能力检测的各项指标对水分胁迫的敏感性顺序为发芽指数>发芽势>发芽率.岷谷木蓝种子的发芽特征是对干旱河谷生存环境长期适应的结果.野外播种的适宜时间为4月初,并在播种前对种子进行7 h的沙摩处理.     

珍稀植物青檀种子休眠与萌发的研究

为了探讨和研究青檀种子休眠和萌发特性,采用石蜡切片法、生物鉴定(白菜籽发芽实验)法、赤霉素溶液浸种、以及赤霉素与低温层积相结合等方法,寻找引起青檀种子休眠的原因和解除休眠的最佳措施。结果表明:青檀种子本身含有发芽抑制物和存在生理后熟是引起休眠的2个主要原因,用质量浓度为300mg/L的赤霉素溶液浸种24h或低温层积后用赤霉素处理均能在一定程度上解除休眠促进萌发,其中以低温层积25d后用500mg/L的赤霉素浸种36h效果最好。发芽率和发芽势分别达到83.5%和65%。     

Seed dormancy and germination in Jeffersonia dubia (Berberidaceae) as affected by temperature and gibberellic acid

The genus Jeffersonia, which contains only two species, has a trans‐Atlantic disjunct distribution. The aims of this study were to determine the requirements for breaking dormancy and germination of J. dubia seeds and to compare its dormancy characteristics with those of the congener in eastern North America. Ripe seeds of J. dubia contain an underdeveloped embryo and were permeable to water. In nature, seeds were dispersed in May, while embryos began to grow in September, and were fully elongated by late November. Germination started in March of the next year, and seeds emerged as seedlings soon after germination. In laboratory experiments, incubation at high temperatures (25 °C, 25/15 °C) for at least 8 weeks was required to initiate embryo growth, while a transfer to moderate temperatures (20/10 °C, 15/6 °C) was needed for the completion of embryo growth. At least 8 weeks at 5 °C was effective in overcoming physiological dormancy and for germination in seeds after the embryos had fully elongated. Thus, both high and low temperatures were essential to break dormancy. Gibberellic acid (GA₃) treatment could substitute for the high temperature requirement, but not for the low temperature requirement. Based on the dormancy‐breaking requirements, it is confirmed that the seeds have deep simple morphophysiological dormancy. This dormancy type is similar to that of seeds of the eastern North American species J. diphylla. Although seeds require 10–11 months from seed dispersal to germination in nature, under controlled conditions they required only 3 months after treatment with 1000 mg·l^?1 GA₃, followed by incubation at 15/6 °C. This represents practical knowledge for propagation of these plants from seed.     

Seed dormancy and germination behaviour in two Euphrasia species (Orobanchaceae) occurring in the Swiss Alps

The optimal time for germination of a seed depends on environmental conditions of its habitat, the life cycle of the germinating plant, and the conditions for successful establishment, growing and reproduction. We studied the germination behaviour of the alpine annual Euphrasia minima and an alpine ecotype of E. salisburgensis in a lowland garden experiment. Seeds of both species and their hybrids germinated at constant (5 °C) and at varying temperatures (3–10 °C), and never before spring after seed ripening. Germination was spread over 3 years, which suggests that the seeds formed a persistent seed bank. The two species together with E. minima and E. minima hybrids differed significantly in the germination rate in the first and second spring.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 649–656.     

Seed dormancy patterns in Epipactis palustris (Orchidaceae): Requirements for germination and establishment of mycorrhiza

Some terrestrial orchid species, including Epipactis palustris (L.) Crantz, are considered extremely difficult to germinate and cultivate in vitro. Observations of orchids germinating in nature are very few, and the timing and requirements for seedling establishment are unknown for most species. Seeds of E. palustris were incubated in vitro with an appropriate fungus, but germination was poor unless several other conditions were also met: scarification of the testa in Ca(OCL)₂, an initial incubation for several weeks at 27°C, and a subsequent cold stratification for 8–12 weeks at 4–8°C, With these pretreatments, germnation responses exceeded 50% after incubation for 4 weeks at 20°C. Healthy protocorms with normal organ development were only produced by symbiotic culture following this lengthy seed preparation. The findings suggest that under natural conditions the seeds need some after-ripening, and the testa needs to be partially decomposed before germination. The requirement for chilling suggests that germination of seeds in situ occurs in spring.     

Seed dormancy in Acer: Endogenous germination inhibitors and dormancy in Acer pseudoplatanus L.

Summary Dormant seeds of Acer pseudoplatanus L. contain two zones of inhibition on paper chromatograms in 10:1:1 as detected by the lettuce and cress seed germination, and the wheat coleoptile bioassays. One zone at R_f 0.6–0.8 was partitioned into ethyl acetate at acid pH and was shown to contain ABA by its behaviour on GLC and isomerization under ultra-violet light. The other zone at R_f 0.9 was detected only in the germination bioassays and was partitioned into ethyl acetate over a range of pH indicating the presence of one or more neutral compounds.The inhibitors present in the embryo of dormant sycamore seeds inhibited the germination of non-dormant sycamore seeds at relatively low concentrations. A comparison with the effects of application of exogenous ABA indicated that endogenous ABA could not solely account for the inhibitory activity of seed extracts, which appeared to be due partly to the presence of ABA and partly to that of neutral compounds present in the embryo. Leaching treatments that removed dormancy led to a decrease in the level of inhibitors present mainly in the basic fraction. The exogenous application of kinetin to dormant sycamore seeds increased germination whereas gibberellic acid had no effect. Similar responses were obtained with lettuce seeds inhibited by the basic fraction of dormant sycamore seeds.It is suggested that an inhibitor-cytokinin interaction may be involved in the dormancy of sycamore seeds.     

Towards a systems biology approach to understanding seed dormancy and germination

Seed germination is the first adaptive decision in the development of many land plants. Advances in genetics and molecular physiology have taught us much about the control of germination using the model plant Arabidopsis thaliana. Here we review the current state of the art with an emphasis on mechanistic considerations and explore the potential impact of a systems biology approach to the problem.     

Seed dormancy and germination in three Crocus ser. Verni species (Iridaceae): implications for evolution of dormancy within the genus

The aim of this work was to examine whether seed ecophysiological traits in three closely related Crocus species were associated with ecological niche differentiation and species divergence. Seeds of the temperate tetraploid cytotype of Crocus neapolitanus, the sub‐Mediterranean C. etruscus and the Mediterranean C. ilvensis were placed either on agar in the laboratory under different periods of simulated seasonal conditions or in nylon mesh bags buried outdoors to examine embryo growth, radicle and shoot emergence. In agreement with the phenology observed outdoors, in the laboratory embryos required a cool temperature (ca. 10 °C) to grow to full size (embryo length:seed length, E:S ratio ca. 0.75) but only after seeds received a warm stratification; radicle emergence then followed immediately (November). Shoot emergence is a temporally separated phase (March) that was promoted by cold stratification in C. neapolitanus while in the other two species this time lag was attributed to a slow continuous developmental process. These species have similar embryo growth and radicle phenology but differ in their degree of epicotyl dormancy, which is related to the length of local winter. Conclusions from laboratory experiments that only consider root emergence could be misleading; evaluating the phenology of both root and shoot emergence should be considered in order to demonstrate ecologically meaningful differences in germination behaviour and to develop effective propagation protocols. Although these taxa resulted from recent speciation processes, the outcomes suggest an early onset of adaptation to local ecological factors and that phylogeny may represent a significant constraint in the evolution and expression of seed traits in Crocus.     

Seed dormancy and control of germination in Sisymbrella dentata (L.) O.E. Schulz (Brassicaceae)

• Seed germination responsiveness to environmental cues is crucial for plant species living in changeable habitats and can vary among populations within the same species as a result of adaptation or modulation to local climates. Here, we investigate the germination response to environmental cues of Sisymbrella dentata (L.) O.E. Schulz, an annual endemic to Sicily living in Mediterranean Temporary Ponds (MTP), a vulnerable ecosystem.
• Germination of the only two known populations, Gurrida and Pantano, was assessed over a broad range of conditions to understand the role of temperatures, nitrate, hormones (abscisic acid – ABA and gibberellins – GA) and after‐ripening in dormancy release in this species.
• Seed germination responsiveness varied between the two populations, with seeds from Gurrida germinating under a narrower range of conditions. Overall, this process in S. dentata consisted of testa and endosperm rupture as two sequential events, influenced by ABA and GA biosynthesis. Nitrate addition caused an earlier testa rupture, after‐ripening broadened the thermal conditions that allow germination, and alternating temperatures significantly promoted germination of non‐after‐ripened seeds.
• Primary dormancy in S. dentata seeds likely allows this plant to form a persistent seed bank that is responsive to specific environmental cues characteristic of MTP habitats.

     

Seed biology and in vitro seed germination of Cypripedium

Abstract

Cypripedium orchids have high horticultural value. The populations of most species are very geographically restricted and they are becoming increasingly rare due to the destruction of native habitats and illegal collection. Reduction of the commercial value through large-scale propagation in vitro is a preferable option to reduce pressure from illegal collection. Cypripedium species are commercially propagated via seed germination in vitro. This review focuses on in vitro seed germination and provides an in-depth analysis of the seed biology of this genus.     

Seed after-ripening and dormancy determine adult life history independently of germination timing

? Seed dormancy can affect life history through its effects on germination time. Here, we investigate its influence on life history beyond the timing of germination. ? We used the response of Arabidopsis thaliana to chilling at the germination and flowering stages to test the following: how seed dormancy affects germination responses to the environment; whether variation in dormancy affects adult phenology independently of germination time; and whether environmental cues experienced by dormant seeds have an effect on adult life history. ? Dormancy conditioned the germination response to low temperatures, such that prolonged periods of chilling induced dormancy in nondormant seeds, but stimulated germination in dormant seeds. The alleviation of dormancy through after-ripening was associated with earlier flowering, independent of germination date. Experimental dormancy manipulations showed that prolonged chilling at the seed stage always induced earlier flowering, regardless of seed dormancy. Surprisingly, this effect of seed chilling on flowering time was observed even when low temperatures did not induce germination. ? In summary, seed dormancy influences flowering time and hence life history independent of its effects on germination timing. We conclude that the seed stage has a pronounced effect on life history, the influence of which goes well beyond the timing of germination.     

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.     

Seed dormancy and germination of five selected NATURA-2000 plant species from Croatia showing different germination strategies

Abstract

NATURA-2000, the ecological network of protected areas in the European Union that has been included in the Croatian legislation, defines Community important plant species with imperative on their conservation ex/in situ. In the Botanical Garden of the Faculty of Science, University of Zagreb, five NATURA-species have been selected as research subjects for germination study to shed light on the topic of their seed ecology and consequently advance their conservation efforts: Degenia velebitica, Scilla litardierei, Klasea lycopifolia, Ligularia sibirica and Genista holopetala. The freshly matured seeds of each species were exposed to cold or warm stratification in duration of four to sixteen weeks, and their germination was investigated through different regimes of incubation parameters, i.e. illumination (light/dark) and temperature (5, 15/6, 23?°C). All species had higher germination values after cold stratification, with the exception of G. holopetala. We concluded that D. velebitica, K. lycopifolia and presumably L. sibirica seeds has non-deep physiological dormancy while S. litardierei has deep complex morphophysiological dormancy and G. holopetala has physical dormancy. The observed patterns in seeds’ behaviour are consistent with the conditions in their natural habitats in Croatia and the knowledge of these patterns is vital for successful conservation strategies in the future.     

Seed germination and dormancy: The classic story,new puzzles,and evolution

This review highlights recent progresses in seed germination and dormancy research.Research on the weakening of the endosperm during germination,which is almost a classic theme in seed biology,was resumed byα-xylosidase studies.Strong genetic evidence was presented to suggest that the quality control of xyloglucan biosynthesis in the endosperm(and the embryo)plays a critical role in germination.Further analyses on the endosperm and the adjace nt layers have suggested that the cutin coat in the endosperm-testa interphase negatively affects germination while the en dosperm-embryo in terphase produces a sheath that facilitates germination.These progresses significantly advanced our understanding of seed germination mechanisms.A breakthrough in dormancy research,on the other hand,revealed the unique abscisic acid signaling pathway that is regulated by DELAY OF GERMINATION1(DOG1).The detailed analysis of DOC1 expression uncovered the intriguing story of reciprocal regulation of the sensean tisense pair,which gen erated new questions.Recent studies also suggested that the DOG1 function is not limited to dormancy but extended through general seed maturation,which provokes questions about the evolution of DOG1 family proteins.Seed biology is becoming more exciting with the classic stories being revitalized and new puzzles emerging from the frontier.