种子休眠机理研究概述

种子休眠是植物本身适应环境和延续生存的一种特性,是种子植物进化的一种稳定对策。野生植物特别是原产温带的植物,其种子大多有深而长的休眠期。关于种子休眠的概念有多种,这些概念引出了许多学说、假说和模型。种壳障碍、胚形态发育不完全和生理后熟以及种子中含有化学抑制剂等,都可导致种子休眠。根据不同的分类标准可将种子分成不同类型,一般将种子分为强迫休眠和机体休眠;机体休眠又可分为外部休眠、内部休眠和综合休眠。植物种类不同休眠特性也不同;同种植物的种子来源于不同的居群和植株时,若采集时期不同,其休眠也可能不同;甚至在同一果实中的不同种子,休眠特性亦可能有差异。影响休眠性状表达的基因既有核基因,也有质基因,休眠通常表现为一种受多基因控制的数量性状。种子休眠具有重要的生态学意义,能有效地调节种子萌发的时空分布。研究种子的休眠特性和机理及其解除方法,有助于农业生产和植物多样性保护。     

Advances in Understanding Seed Dormancy at the Whole-seed Level:An Ecological, Biogeographical and Phylogenetic Perspective

Following a brief account of the early foundations of seed germination ecology sensu lato, some historical and recent developments pertaining to the ecology, biogeography and phylogeny of seed dormancy are discussed.     

植物种子休眠的原因及休眠的多形性

概述了植物种子休眠的原因及种子休眠的多形性。种壳障碍、胚形态发育不全和生理后熟以及种子中含有化学抑制剂都可导致种子休眠。根据不同的分类标准可将种子休眠可分成不同类型,但通常将休眠分为外源休眠、内源休眠和综合休眠。影响休眠的因素是复杂的,植物种类不同,休眠特性不同;同种植物的种子,来源于不同居群和植株,在不同时期采集,位于母株不同位置,其休眠有可能不同;甚至同一果实中的不同种子,休眠特性都会有差异。影响休眠性状表达的基因既有核基因,又有质基因,休眠通常表现为一种受多基因控制的数量性状。种子休眠的多形性有利于调节种子萌发的时空分布。     

Diversity of epicotyl dormancy among tropical montane forest species in Sri Lanka

• Fruiting season of many Sri Lankan tropical montane species is not synchronised and may not occur when conditions are favourable for seedling establishment. We hypothesised that species with different fruiting seasons have different seed dormancy mechanisms to synchronise timing of germination with a favourable season for establishment. Using six species with different fruiting seasons, we tested this hypothesis.
• Germination and imbibition of intact and manually scarified seeds were studied. Effect of GA₃ on germination was examined. Embryo length:seed length (E:S) ratio of freshly matured seeds and of those with a split seed coat was determined. Time taken for radicle and plumule emergence and morphological changes of the embryos were recorded.
• The radicle emerged from Ardisia missionis, Bheza nitidissima and Gaetnera walkeri seeds within 30 days, whereas it took >30 days in other species. Embryos grew in seeds of B. nitidissima and G. walkeri prior to radicle emergence but not in Microtropis wallichiana, Nothapodytes nimmoniana and Symplocos cochinchinensis. A considerable delay was observed between radicle and plumule emergence in all six species. Warm stratification and/or GA₃ promoted germination of all species.
• All the tested species have epicotyl dormancy. Seeds of B. nitidissima and G. walkeri have non‐deep simple morphophysiological epicotyl dormancy, and the other four species have non‐deep physiological epicotyl dormancy. Differences in radicle and epicotyl dormancy promote synchronisation of germination to a favourable time for seedling development. Therefore, information on dormancy‐breaking and germination requirements of both radicle and epicotyl are needed to determine the kind of dormancy of a particular species.

     

羊草种子休眠机制及破除方法研究

羊草种子休眠程度深、发芽率低是限制栽培利用的重要因子.采用不同破除羊草种子休眠的方法,测定各处理对种子萌发的影响,以探索破除羊草种子休眠的有效途径.结果显示:(1)刺破种皮的裸种子较完整种子的萌发率、吸水速率、生活力分别由对照的6%、63%、0%显著增加到60%、86%、94%.(2)完整羊草种子分别用清水浸种1 d、30% NaOH浸种80 min、清水浸种1 d后用30% NaOH浸种60min其萌发率由6%分别显著提高到36%、60%、84%,而各浓度赤霉素处理完整种子其萌发率较对照均无显著变化. (3)采用清水浸种1 d后用30% NaOH处理60 min,再施加200 μg/g GA3综合处理,可使羊草完整种子的发芽率由6%提高到91%,接近其种子生活力94%.研究表明,羊草种子的稃与种皮不影响种子水分的吸收,但影响种子对GA3的吸收、不同程度地阻碍大分子物质的渗入、限制羊草种子内部萌发抑制物的渗出,从而引起种子休眠;分析认为稃和种皮以及种子内部萌发抑制物质是引起羊草种子休眠的主要原因.     

Dormancy and germination: making every seed count in restoration

From 50 to 90% of wild plant species worldwide produce seeds that are dormant upon maturity, with specific dormancy traits driven by species' occurrence geography, growth form, and genetic factors. While dormancy is a beneficial adaptation for intact natural systems, it can limit plant recruitment in restoration scenarios because seeds may take several seasons to lose dormancy and consequently show low or erratic germination. During this time, seed predation, weed competition, soil erosion, and seed viability loss can lead to plant re‐establishment failure. Understanding and considering seed dormancy and germination traits in restoration planning are thus critical to ensuring effective seed management and seed use efficiency. There are five known dormancy classes (physiological, physical, combinational, morphological, and morphophysiological), each requiring specific cues to alleviate dormancy and enable germination. The dormancy status of a seed can be determined through a series of simple steps that account for initial seed quality and assess germination across a range of environmental conditions. In this article, we outline the steps of the dormancy classification process and the various corresponding methodologies for ex situ dormancy alleviation. We also highlight the importance of record‐keeping and reporting of seed accession information (e.g. geographic coordinates of the seed collection location, cleaning and quality information, storage conditions, and dormancy testing data) to ensure that these factors are adequately considered in restoration planning.     

Combinational dormancy in seeds of Sicyos angulatus (Cucurbitaceae,tribe Sicyeae)

Seeds with a water‐impermeable seed coat and a physiologically dormant embryo are classified as having combinational dormancy. Seeds of Sicyos angulatus (burcucumber) have been clearly shown to have a water‐impermeable seed coat (physical dormancy [PY]). The primary aim of the present study was to confirm (or not) that physiological dormancy (PD) is also present in seeds of S. angulatus. The highest germination of scarified fresh (38%) and 3‐month dry‐stored (36%) seeds occurred at 35/20°C. The rate (speed) of germination was faster in scarified dry‐stored seeds than in scarified fresh seeds. Removal of the seed coat, but leaving the membrane surrounding the embryo intact, increased germination of both fresh and dry‐stored seeds to > 85% at 35/20°C. Germination (80–100%) of excised embryos (both seed coat and membrane removed) occurred at 15/6, 25/15 and 35/20°C and reached 95–100% after 4 days of incubation at 25/15 and 35/20°C. Dry storage (after‐ripening) caused an increase in the germination percentage of scarified and of decoated seeds at 25/15°C and in both germination percentage and rate of excised embryos at 15/6°C. Eight weeks of cold stratification resulted in a significant increase in the germination of scarified seeds at 25/15 and 35/20°C and of decoated seeds at 15/6 and 25/15°C. Based on the results of our study and on information reported in the literature, we conclude that seeds of S. angulatus not only have PY, but also non‐deep PD, that is, combinational dormancy (PY + PD).     

种子休眠与破眠机理研究进展

种子休眠机理主要围绕透性、抑制剂作用和光敏素转化等方面的研究而建立。种皮的阻碍作用可能是由于种皮的物理或化学特性引起．可导致对水、光、气体或溶质的透性改变。抑制剂作用机理是抑制物质可抵消促进细胞分裂和生长发育的激素的作用。光敏素转化机理来源于与休眠有关的生物活性化学物质的合成、活化或破坏受光诱导的观点,由于发现了光敏素蓝色蛋白的活化型（Pfr）和钝化型（Pr）而得到强有力的支持,种子光休眠取决于光敏素蓝色蛋白的活化型（Pfr）含量和Pfr／（Pr＋Pfr）比值。目前,打破休眠的方法一般有机械破皮法、激素处理法、分子生物学技术法、物理处理法（如激光、烟、热等处理技术）、CO2处理法等。激素的平衡由抑制剂占优势向促进物占优势的变化是打破休眠的决定因素。研究破眠机理的分子生物学技术有多种,包括ABA突变体的利用、分子标记、转基因技术、用反义RAN阻止基因的表达、cDNA克隆技术等。用激光照射种子,把适宜的光射入细胞,可增加细胞生物能,促进种子发育,从而可能打破休眠。热处理的机理是由于加热可以增加种皮的透气性。CO2之所以能提高某些物种的萌发率,在于其影响了种子内部乙烯的敏感性。     

Advances in Understanding Seed Dormancy at the Whole-seed Level: An Ecological,Biogeographical and Phylogenetic Perspective

Following a brief account of the early foundations of seed germination ecology sensu lato , some historical and recent developments pertaining to the ecology, biogeography and phylogeny of seed dormancy are discussed .     

Effect of Temperature Regimes on Germination of Dimorphic Seeds of Atriplex prostrata

Dimorphic seeds of Atriplex prostrata were removed from cold dry storage monthly over a one year period to test for fluctuations in seed dormancy and germination rate. For each seed type, four replicates of 25 seeds were exposed to four alternating night/day temperature regimes mimicking seasonal fluctuations in Ohio: 5/15 °C; 5/25 °C; 15/25 °C and 20/35 °C with a corresponding 12-h photoperiod (20 μmol m⁻² s⁻¹; 400 – 700 nm). We found a significant three-way interaction of seed size, temperature and month for both percent germination and the rate of germination. Large seeds showed the greatest germination at the 20/35 °C and 5/25 °C temperature regimes and small seeds at the 5/25 °C regime. Large seeds had greater germination at all temperatures as compared to small seeds. Large seeds had the fastest germination rates at 20/35 °C followed by 5/25 °C whereas small seeds had the fastest rates at 5/25 °C followed by 20/35 °C. This revised version was published online in July 2006 with corrections to the Cover Date.     

药用植物种子休眠类型及休眠解除研究进展

种子休眠是植物在长期发育过程中形成的对不良环境条件主动适应的现象。介绍了常见药用植物种子休眠的原因,根据休眠原因对种子休眠与萌发的影响将种子休眠划分的几种类型,总结了对不同休眠类型的种子应分别采用的休眠解除方法,并评述了目前的研究进展。     

不同种子预处理对10种沙拐枣植物萌发的影响

为了确定沙拐枣植物种子的萌发特性及最优播前预处理方法,在实验室条件下,对10种沙拐枣植物的种子进行了磨砺、硫酸和热水浸泡、冷藏、种子浸出液处理,然后进行发芽实验研究。萌发实验的结果表明,10种沙拐枣植物对于不同的种子预处理,均表现出相似的萌发反应。磨砺、硫酸浸泡和冷藏处理对种子萌发有明显地促进作用。与对照相比,种子浸出液处理对种子的发芽率、发芽速度均具有明显地抑制作用,并能增强种子的休眠。冷藏处理具有打破有活力的种子休眠、促进种子萌发的作用,但它与热水浸泡处理一样,对有活力种子表现出一定的致死作用。沙拐枣植物的萌发模式在不同种子预处理问均表现出明显的差异性。机械磨擦和硫酸处理能够促进种子的萌发率及发芽势。泡果沙拐枣(Callingonum junceum)在本项实验中表现出很强的萌发能力。     

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。     

脱落酸调控种子休眠和萌发的分子机制

脱落酸(ABA)是调控种子休眠和萌发过程的主要植物激素。种子内源ABA含量和种胚对ABA敏感性共同调控种子休眠和萌发过程, 确保植物种子以休眠状态在逆境中保持其自身繁衍能力, 并在适宜的环境下启动萌发程序。种子ABA合成代谢和ABA信号转导途径涉及许多重要基因家族, 它们通过复杂的调控网络精确地控制着种胚发生、种子成熟、休眠及萌发进程。该文对ABA调控种子休眠和萌发的分子机制最新研究进展进行综述, 并展望了今后的研究方向。     

Morphophysiological dormancy in seeds of the ANA grade angiosperm Schisandra arisanensis (Schisandraceae)

We determined the kind of seed dormancy in Schisandra arisanensis, an ANA grade ([A]mborellales [N]ymphaeales [A]ustrobaileyales) angiosperm with medicinal value. Seeds have small underdeveloped embryos, and following seed maturity their length increased approximately 360% before radicle emergence. Germination was delayed 6–8 weeks, and the percentage and rate were much higher at 15/6, 20/10 and 25/15°C than at 30/20°C. For seeds incubated at 5/5°C (8 weeks) → 15/6°C (4 weeks) → 20/10°C (8 weeks) → 25/15°C (12 weeks) → 20/10°C (5 weeks), embryos grew at 15/6°C → 20/10°C, and almost all seeds that germinated (89%) did so at 20/10°C → 25/15°C. When seeds were incubated in a complementary temperature sequence, 25/15°C (12 weeks) → 20/10°C (8 weeks) → 15/6°C (4 weeks) → 5/5°C (9 weeks) → 15/6°C (4 weeks), embryos grew at 25/15°C → 20/10°C. Nearly all seeds that germinated (93%) did so at 25/15°C → 20/10°C and at 15/6°C following 9 weeks at 5/5°C. Based on the temperature requirements for embryo growth and seed germination, seeds of this species have non‐deep simple morphophysiological dormancy (C_1bB).     

风箱果种子催芽技术研究

以风箱果自然种群成熟的种子为研究对象,对种子的生活力和催芽方法进行了研究,结果表明：TTC染色法测得风箱果种子的生活力达到94.44%;清水浸种处理的发芽势和发芽率均较低,以浸种48 h的效果较好,发芽率为28.33%;不同化学药剂处理均能不同程度的提高发芽率,其中50 mg·L^-1 GA₃+4%KNO₃的混合药剂处理72 h的萌发效果最好,发芽率达到90%;100和200 mg·L^-1 GA₃能极显著提高发芽势（P<0.01）;低温处理28 d能显著提高发芽势和发芽率（P<0.05）。     

Seed survival of the tropical tree Cryptocarya aschersoniana (Lauraceae): Consequences of habitat disturbance

Seed survival in soil could be strongly influenced by habitat characteristics, but little is known about the behaviour of seeds sensitive to desiccation in seed banks installed in natural or disturbed habitats. Cryptocarya aschersoniana seeds disperse at the end of the rainy season but do not germinate immediately; thus, they may form seed banks in soil. This study evaluated the behaviour of C. aschersoniana seed banks induced in the natural environment of the species and in a disturbed area. Recently harvested C. aschersoniana germination units were characterized according to their water content, germination and viability. In 2011 and 2012, seed banks were established by burying samples of seeds in the understory of a semi‐deciduous forest. In 2012, samples were also buried in a disturbed area. The seed banks were sampled at certain time intervals, and the samples were characterized as described above. Precipitation and air temperature data were collected. As a result, seeds in the seed bank established in the natural environment form a transient seed bank and showed the same behaviour in both years studied. A germination peak was observed starting 210 days after burial (coinciding with the onset of the rainy season) and reached germination percentages higher than 80% at the end of the experiment for both years. Seed mortality did not exceeded 28% in the natural environment. However, in the disturbed environment, the seeds lost their viability more rapidly, with 90% of the seeds becoming unviable 240 days after burial. Germinated seeds in the disturbed environment (maximum 21%) were not able to establish seedlings. These results underscore the importance of maintaining a natural, undisturbed forest for the conservation of this species.     

苦槠传播体的萌发特性与种子休眠类型

种子的萌发特性是植物在长期进化过程中形成的一种适应环境变化的固有属性。对种子萌发特性的研究有助于我们理解物种的种群更新和种子育苗。本文旨在阐明我国重要资源植物苦槠（Castanopsis sclerophylla）的种子休眠类型和萌发特性。研究结果发现：①苦槠种子在其果实中占比非常高;②苦槠的果皮限制了种子对水分的吸收;③苦槠果皮阻碍了种子的萌发,除去果皮以后种子能够在较大温度范围内萌发;④除去果皮以后,苦槠种子在低温（15/5 ℃、20/10 ℃）下萌发困难,在25/15 ℃、30/20 ℃和35/25 ℃这3个温度梯度下均能迅速萌发。结果表明,苦槠的传播体（果实）具有物理休眠的特性。因此,壳斗科（Fagaceae）是具有物理休眠和脱水敏感性的一个新纪录科。结合壳斗科的系统发育位置,本研究支持物理休眠是一种较为进化的性状这一假说。     

Dormancy cycles in buried seeds of three perennial Xyris (Xyridaceae) species from the Brazilian campo rupestre

• Dormancy cycles are an important mechanism for avoiding seed germination under unfavourable periods for seedling establishment. This mechanism has been scarcely studied in tropical species. Here, we studied three tropical and perennial species of Xyris, X. asperula, X. subsetigera and X. trachyphylla, to investigate in situ longevity and the existence of seasonal seed dormancy cycles.
• Seeds of three species of Xyris were buried in their natural habitat, with samples exhumed bimonthly for 18 months. Germination of exhumed seeds was assessed under a 12‐h photoperiod over a broad range of temperatures. Seeds of X. trachyphylla were also subjected to treatments to overcome secondary dormancy.
• Seeds of all species are able to form a persistent seed bank and exhibit seasonal changes in germinability. Secondary dormancy was acquired during the rainy summer and was overcome during the subsequent dry season (autumn/winter). Desiccation partially overcomes secondary dormancy in X. trachyphylla seeds.
• Soil seed bank persistence and synchronisation of seed germination under favourable conditions for seedling establishment contribute to the persistence and regeneration of X. asperula, X. subsetigera and X. trachyphylla in their natural environment.

     

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.