Investigating seed dormancy in cotton (Gossypium hirsutum L.): understanding the physiological changes in embryo during after‐ripening and germination

• The dormancy of seeds of upland cotton can be broken during dry after‐ripening, but the mechanism of its dormancy release remains unclear.
• Freshly harvested cotton seeds were subjected to after‐ripening for 180 days. Cotton seeds from different days of after‐ripening (DAR) were sampled for dynamic physiological determination and germination tests. The intact seeds and isolated embryos were germinated to assess effects of the seed coat on embryo germination. Content of H₂O₂ and phytohormones and activities of antioxidant enzymes and glucose‐6‐phosphate dehydrogenase were measured during after‐ripening and germination.
• Germination of intact seeds increased from 7% upon harvest to 96% at 30 DAR, while embryo germination improved from an initial rate of 82% to 100% after 14 DAR. Based on T₅₀ (time when 50% of seeds germinate) and germination index, the intact seed and isolated embryo needed 30 and 21 DAR, respectively, to acquire relatively stable germination. The content of H₂O₂ increased during after‐ripening and continued to increase within the first few hours of imbibition, along with a decrease in abscisic acid (ABA) content. A noticeable increase was observed in gibberellic acid content during germination when ABA content decreased to a lower level. Coat removal treatment accelerated embryo absorption of water, which further improved the accumulation of H₂O₂ and changed peroxidase content during germination.
• For cotton seed, the alleviation of coat‐imposed dormancy required 30 days of after‐ripening, accompanied by rapid dormancy release (within 21 DAR) in naked embryos. H₂O₂ acted as a core link between the response to environmental changes and induction of other physiological changes for breaking seed dormancy.

     

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.     

Morpho‐colorimetric analysis and seed germination of Brassica insularis Moris (Brassicaceae) populations

Brassica insularis is a perennial plant growing on both coastal and inland cliffs. Three seed lots from Sardinia were analysed using an image analysis system to detect differences in seed morphology, both within and among populations. Germination requirements at constant (5–25 °C) and alternating temperatures (25/10 °C), both in light and in darkness, were evaluated for all populations. In addition, the effect of a dry after‐ripening period (90 days at 25 °C) was also investigated. Morpho‐colorimetric analysis clearly identified seeds from different populations and discriminated three chromatic categories for seeds belonging to the Isola dei Cavoli coastal population, but not for the inland Masùa and the coastal Planu Sartu. Inter‐population variability was also observed in germination behaviour. B. insularis seeds germinated, with percentages up to 60%, in a wide range of temperatures (5–25 °C), and neither light nor dry after‐ripening affected final germination percentages. Moisture content measurements were made for seeds of each colour, but there were no particular differences among colours. Inter‐populational variability in germination behaviour may be a survival strategy for species growing under unpredictable environmental conditions, such as under Mediterranean climate, while heteromorphy may be due to independent evolutionary divergence processes of the Isola dei Cavoli population.     

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 traits and germination behaviour of four Sardinian populations of Helichrysum microphyllum subsp. tyrrhenicum (Asteraceae) along an altitudinal gradient

• Helichrysum microphyllum subsp. tyrrhenicum (Asteraceae) is an endemic taxon of Sardinia and Corsica, where it grows at different altitudes. The objective of this study was to investigate the seed traits and germination behaviour of four Sardinian populations of this taxon located at different altitudes.
• Seed traits were evaluated, and germination tests were carried out by incubating seeds at a range of constant (5–30 °C) and alternating (25/10 °C) temperatures. The dry after‐ripening (DAR) pre‐treatment was also applied by storing seed in dry conditions for 3 months at 25 °C. Seed traits and germination behaviour data were statistically analysed to identify if there was a correlation with altitude.
• Differences in seed size, area and mass among populations were recorded, however, no relationship was found with altitude. High germination percentages were obtained in all populations, both in untreated and DAR seeds, and were positively affected by alternating temperatures. The final germination percentage and time required to reach 50% final germination (T₅₀) showed no relationship with altitude.
• The differences in seed traits and germination detected among the studied populations of H. microphyllum subsp. tyrrhenicum were not correlated with altitude. This study provides new and important knowledge for this taxon. H. microphyllum subsp. tyrrhenicum is characterised by high germination percentages and low T₅₀ values and does not seem to require any dormancy‐breaking treatment. This species represents a high‐potential native plant species that should be considered within environmental management plans.

     

Seed dormancy of Cardiospermum halicacabum (Sapindaceae) from three precipitation zones in Sri Lanka

• This study investigated seed germination of Cardiospermum halicacabum, a medicinally important invasive species.
• We compared mass, moisture content (MC), dormancy and dormancy‐breaking treatments and imbibition and germination of scarified and non‐scarified seeds of C. halicacabum from a low‐elevation dry zone (DZ), low‐elevation wet zone (WZ1) and mid‐elevation wet zone (WZ2) in Sri Lanka to test the hypothesis that the percentage of seeds with water‐impermeable seed coats (physical dormancy, PY) decreases with increased precipitation.
• Seed mass was higher in WZ2 than in DZ and WZ1, while seed MC did not vary among the zones. All scarified DZ, WZ1 and WZ2 and non‐scarified DZ and WZ1 seeds imbibed water, but only a few non‐scarified WZ2 seeds did so. When DZ and WZ1 seeds were desiccated, MC and percentage imbibition decreased, showing that these seeds have the ability to develop PY. GA₃ promoted germination of embryos excised from fresh DZ and WZ1 seeds and of scarified WZ2 seeds.
• At maturity, seeds from DZ and WZ1 had only physiological dormancy (PD), while those from WZ2 had combinational dormancy (PY+PD). Thus, our hypothesis was not supported. Since a high percentage of excised embryos developed into normal seedlings; this is a low‐cost method to produce C. halicacabum plants for medicinal and ornamental purposes.

     

Differences in seed dormancy and germination in amphicarpic legumes: manifold bet-hedging in space and time

两型结实豆科植物种子休眠和萌发的差异：空间和时间上多样性的两头下注策略通过休眠而实现的在空间或时间上的传播是生物体对它们所经历的环境施加影响的主要过程之一。在植物中,由于这两种传播都是通过种子进行的,而且两者在适应性方面发挥的作用也是类似的,因此预测它们之间存在较强的进化相关性。在本研究中,我们使用了两型结实的植株来研究这些进化相关性,这类植株能同时产生具有高度空间传播力的种子以及不会传播的地下种子。我们对经不同组合的休眠过程破坏处理(即干燥后熟、冷积层和物理划伤)后的种子的萌发百分比进行了评估,以研究两种两型结实豆科植物——野豌豆(Vicia amphicarpa L.)和山黧豆(Lathyrus amphicarpos L.),以及同它们亲缘关系很近的两种单型结实类群——救荒野豌豆(Vicia sativa L.)和红山黧豆(Lathyrus cicera L.)的飞播种子和地下种子在休眠和萌发方面的变化情况。研究结果表明,空间传播和时间传播之间存在着复杂的相互作用。在刚完成传播时,空播种子比地下种子更具休眠性,但当种子经后熟处理而开始物理休眠时,这种趋势也随之逆转。单型结实植株种子的萌发百分比高于其两型结实的同属植株,并在后熟处理后都同样失去了休眠性。相反,两型结实植株的种子则表现出同时受生理休眠和物理休眠调节的不同休眠性策略,预计这可能使出苗时间发生变化,从而提供了多层次多样化的两头下注策略(bet-hedging)。这一策略能够使植株依赖于丰产季中的历史有利区域而不阻碍空间和时间上的迁移,从而有可能在不可预测性很高的条件下具有适应性。     

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。     

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。     

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.     

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.     

Temperature conditions control embryo growth and seed germination of Corydalis solida (L.) Clairv., a temperate forest spring geophyte

Spring is often the most suitable period for seedling establishment of temperate woodland species. Different physiological mechanisms resulting in spring emergence have evolved in seeds of such plants. The aim of this study was to determine the requirements for breaking dormancy and for seed germination of the European perennial spring geophyte Corydalis solida (Fumariaceae). Ripe seeds of C. solida contain an underdeveloped embryo, consisting of no more than a clump of cells. As a consequence, the embryo has to differentiate and grow to a critical length before germination can occur. In nature, seeds are dispersed in spring, while growth of the embryo starts in the autumn and continues in winter. Germination starts in late winter, immediately after embryo growth is completed, resulting in seedling emergence in the following spring. Experiments in controlled conditions showed that temperature is the main factor controlling dormancy and germination. Incubation at autumn temperatures (15/6 °C; 20/10 °C) for at least 8 weeks is required to initiate embryo growth, while a transfer to 5 °C is needed for completion of embryo growth and germination. Growth of the embryo of C. solida occurs at different temperatures over an extended period, a feature typical of temperate forest herbs. Our results indicate that the dormancy mechanism in seeds of C. solida is very similar to mechanisms in other Corydalis species studied thus far, suggesting that stasis in the dormancy trait has occurred.     

Seed germination and dormancy in the medicinal woodland herbs Collinsonia canadensis L. (Lamiaceae) and Dioscorea villosa L. (Dioscoreaceae)

We used a double germination phenology or “move-along” experiment (sensu Baskin and Baskin, 2003) to characterize seed dormancy in two medicinal woodland herbs, Collinsonia canadensis L. (Lamiaceae) and Dioscorea villosa L. (Dioscoreaceae). Imbibed seeds of both species were moved through the following two sequences of simulated thermoperiods: (a) 30/15 °C→20/10 °C→15/6 °C→5 °C→15/6 °C→20/10 °C→30/15 °C, and (b) 5 °C→15/6 °C→20/10 °C→30/15 °C→20/10 °C→15/6 °C→5 °C. In each sequence, seeds of both species germinated to high rates (>85%) at cool temperatures (15/6 and 20/10 °C) only if seeds were previously exposed to cold temperatures (5 °C). Seeds kept at four control thermoperiods (5, 15/6, 20/10, 30/15 °C) for 30 d showed little or no germination. Seeds of both species, therefore, have physiological dormancy that is broken by 12 weeks of cold (5 °C) stratification. Morphological studies indicated that embryos of C. canadensis have “investing” embryos at maturity (morphological dormancy absent), whereas embryos of D. villosa are undeveloped at maturity (morphological dormancy present). Because warm temperatures are required for embryo growth and cold stratification breaks physiological dormancy, D. villosa seeds have non-deep simple morphophysiological dormancy (MPD). Neither species afterripened in a 6-month dry storage treatment. Cold stratification treatments of 4 and 8 weeks alleviated dormancy in both species but C. canadensis seeds germinated at slower speeds and lower rates compared to seeds given 12 weeks of cold stratification. In their natural habitat, both species disperse seeds in mid- to late autumn and germinate in the spring after cold winter temperatures alleviate endogenous dormancy.     

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.     

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.     

外来杂草小花山桃草种子休眠萌发特性

种子萌发和出苗是植物一生中对环境胁迫最为敏感的阶段,外来植物在这一时期对环境条件的适应能力是决定其入侵能力大小的关键因素之一。通过室内外试验研究了小花山桃草种子休眠特性以及贮藏时间、果皮、温度、p H值、水势、盐分和埋土深度对其种子萌发、幼苗生长和出苗的影响。结果表明:小花山桃草蒴果中4粒种子的大小差异显著(P0.01),但它们的活力和萌发率差别不大,4粒种子活力均高达96%以上。休眠解除后,萌发率均高达98%以上;坚硬果皮的机械阻碍是引起种子休眠的主要原因;在室温储藏条件下,种子萌发率随贮藏时间的增加先增加后降低。贮藏1个月,萌发率提高到54%(贮藏前为0)。贮藏3—6个月时,萌发率达最大值(98.5%—99%)。贮藏1a后,萌发率和萌发速度显著下降,但萌发率仍高达90%以上;在室外,有10.5%的蒴果前两粒种子(果实柱头端)同时萌发出苗。室外埋藏1a后,蒴果中未萌发的3粒种子的发芽率分别为45.5%、90.5%和88.5%;小花山桃草种子萌发的温度范围为5—35℃,15—25℃最适于种子萌发,25℃最有利于幼苗生长;小花山桃草能忍受一定的盐胁迫和干旱胁迫,盐浓度和水势为0.15 mol/L和-0.5 MPa时,种子萌发率分别为33.5%和9%。种子萌发和幼苗生长最适宜的土壤含水量为50%;小花山桃草对p H值具有较广泛的适应性,在p H值为5—9范围内均可萌发;小花山桃草种子在土壤表面的出苗率为43%,埋土深度2 cm时,出苗率最高(89.5%),埋土深度大于10 cm时,不再出苗。小花山桃草种子休眠萌发特征以及出苗特点是其快速入侵的基础。研究结果为预测小花山桃草潜在分布范围以及提出科学合理的综合治理措施提供了理论依据。