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 .     

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。     

红松种子休眠与种皮的关系

本文探讨红松(Pinus koraiensis)种子休眠与其种皮之间的关系。夹破中种皮后,种子萌发率很低。在离体胚培养基中外加 ABA 及经 ABA 溶液浸泡种子的萌发实验表明,ABA也不是导致休眠的关键因素。试验确认红松种子存在透气障碍,即中、内种皮对氧气的进入都有阻碍作用。经低温砂藏后,种皮的阻碍作用明显减小。种皮的透气性障碍可能是诱导休限的主导因素。     

Seed germination ecology of Polymnia canadensis (Asteraceae), a monocarpic species of the North American Temperate Deciduous Forest

The seed germination of Polymnia canadensis was studied in relation to its wide variation in life history and its ecology as a mostly facultative biennial, a life cycle type that is rare in stable forest habitats in eastern North America. The seed dormancy and germination characteristics of P. canadensis were like some other herbaceous woodland species of diverse life cycle types. That is, its seeds have physiological dormancy, and they (1) germinated to higher percentages after cold stratification, (2) germinated better in light than in darkness and (3) formed a persistent soil seed bank. Since its seed dormancy and germination characteristics are not uncommon and are not particularly associated with any one life cycle type in woodlands, those properties do not contribute to the rarity of its life cycle type in deciduous forests. These properties are also found in many facultative biennials of open, disturbed habitats and in other Asteraceae of various life cycle types. In addition, the germination phenology of P. canadensis differed between mesic and dry field sites in central Kentucky. At the dry sites, most of the yearly germination occurred in fall, while at the mesic sites most germination occurred in the spring due to being inhibited during fall. A reciprocal seed transplant experiment conducted in two different years demonstrated that the cause of this difference was environmental and that there was no source x site interaction; i.e., no difference between sources in phenotypic plasticity for germination phenology. Absence of genetic effects was also shown by no consistent pattern in the few differences between seed sources in seed incubation laboratory experiments and in a germination phenology experiment in glasshouse flats. Exposure of seeds to a high ratio of far-red/red light prior to or during incubation did not inhibit germination. Neither did changes in leaf litter thickness qualitatively alter germination phenology in field quadrats or in glasshouse flats. Thus, germination in fall at mesic sites was inhibited by other unknown factor(s), perhaps interacting with the effects of closed tree canopy and/or thick leaf litter.     

R.M. Cowling,D.M. Richardson & S.M. Pierce (eds.) 1977. Vegetation of Southern Africa.

We determined the role of bird dispersal in seed and seedling dynamics of the tree Kalopanax pictus from 7 years of observing seed rain and seedling emergence in a broad-leaved deciduous forest in central Japan. We also performed an experiment on the influence of seed pulp on germination of seeds of K. pictus. Seeds of this species can lie dormant for several years, and this causes rather constant yearly seedling emergence in spite of irregular seed production. The spatial distribution of the seedlings that emerged each year (maximum distance from nearest conspecific seed-bearing tree of 90 m) was wider than that of gravity-dispersed seeds (max. distance of 37 m), suggesting seed dispersal by birds in winter. Emerged seedling densities at sites over 20 m from the nearest conspecific seed-bearing tree were highest in the spring of 1991, about half a year after the largest seed fall of the observation period. However, emerged seedling densities within 20 m from seed-bearing trees were highest in 1992, 1.5 years after the largest seed fall. These field observations may be explained by the experimental results on the effects of seed pulp on germination. Intact seeds germinate slowly at low germination rates, while seeds without seed pulp germinate quickly at high germination rates. Fallen seeds with seed pulp thus appear to form a seed bank near seed sources (temporal dispersal), while seeds scattered by birds appear to increase the possibility of reaching the present safe sites in distant areas with quick germination (spatial dispersal).     

白皮松种子休眠特性的种源变异

对７个具有代表性种源的白皮松种子的休眠特性研究表明,白皮松种子种皮的透水性在种源间差异明显,表现出随地理纬度的增高而逐渐增大的趋势,分布越是偏北的种源透水性趋于加强。根据种皮的透水性,７个种源可被分为差异明显的２组,组间以秦岭为界。不同种源种子种皮的透气性以外种皮造成的差异为最大,内种皮次之。种子的透气性与透水性表现出的变化趋势相同。不同种源种子浸提液的抑制能力显著不同,而且这种差异随浸提液浓度的     

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.     

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.     

Climate change and plant regeneration from seed

At the core of plant regeneration, temperature and water supply are critical drivers for seed dormancy (initiation, break) and germination. Hence, global climate change is altering these environmental cues and will preclude, delay, or enhance regeneration from seeds, as already documented in some cases. Along with compromised seedling emergence and vigour, shifts in germination phenology will influence population dynamics, and thus, species composition and diversity of communities. Altered seed maturation (including consequences for dispersal) and seed mass will have ramifications on life history traits of plants. Predicted changes in temperature and precipitation, and thus in soil moisture, will affect many components of seed persistence in soil, e.g. seed longevity, dormancy release and germination, and soil pathogen activity. More/less equitable climate will alter geographic distribution for species, but restricted migratory capacity in some will greatly limit their response. Seed traits for weedy species could evolve relatively quickly to keep pace with climate change enhancing their negative environmental and economic impact. Thus, increased research in understudied ecosystems, on key issues related to seed ecology, and on evolution of seed traits in nonweedy species is needed to more fully comprehend and plan for plant responses to global warming.     

Seed size and seed germination in the Mediterranean fire-prone shrub <Emphasis Type="Italic">Cistus </Emphasis><Emphasis Type="Italic">ladanifer</Emphasis>

The germination response of different sized seeds from individuals of a Mediterranean fire-prone shrub (Cistus ladanifer) was investigated in relation to pre-germination heating. A control (no heating), a low temperature during a short exposure time (50°C during 5 min), a high temperature during a short exposure time (100°C during 5 min) and a high temperature during a long exposure time (100°C during 15 min) were applied to seeds from different individual plants with different mean seed weight. These pre-germination treatments resemble natural germination scenarios for the studied species, absence of fire, low intensity pasture fire, typical Mediterranean shrub fire, and severe fire with high fuel load. Mean seed weight only showed a marginally significant positive correlation with the proportion of germinated seeds whatever the pre-germination treatment. These results suggest that seed dormancy is unrelated to seed size and that under the experimental conditions used in this study, the effect of seed size on seed germination is low. Nevertheless, larger seeds could be favoured in natural conditions, especially under the high competition scenario which arise after wildfires. Control seeds showed a negative correlation between seed size and germination velocity suggesting that lighter seeds could take advantage from early germination in recruitment events in the absence of wildfires. Nevertheless, even the lower pre-germination heating treatment turns this correlation in not significant, suggesting a strong selection pressure (unrelated to seed size) for early germination after fire events. In our study, different sized seeds of C. ladanifer seem to perform better under different germination scenarios suggesting that seed size variation could be maintained by the alternation of recruitments without wildfires and recruitments after wildfire events.     

Structural Features Underlying Hardseededness in Geraniaceae

Abstract: The proportion of hard Geraniaceae seeds ranges from 0 to 100%, depending on species. By analysing the characteristics of hardseededness through the pathway of water uptake in soft seeds provided localization of the chalazal area in water uptake. This represents the most important feature of seed coat permeability. The structural difference between hard-and softseededness was clarified by comparing different species with exclusively permeable or impermeable seeds. Soft seeds form a wide opening at the chalazal slit, while hard ones close the slit using adjacent palisade cells so effectively continuing the impermeable barrier.     

鸢尾属植物种子休眠研究进展

鸢尾属(Iris L.)花卉具有较高的观赏性,极具开发利用价值,且主要以种子繁殖后代;但多数种子具有休眠特性,给育种工作带来了困难。其休眠原因包括种皮机械阻碍和种子中萌发抑制物存在等内外多重因素,单一或交互作用影响种子萌发。种子休眠解除的方法主要有环境条件处理、机械处理、化学试剂或激素处理等。本文对鸢尾属植物种子休眠原因、休眠解除方法及其机理等作了系统概述,旨在为鸢尾属花卉资源的合理开发利用及种质创新提供理论依据。     

松嫩碱化草甸土壤种子库格局、动态研究进展

草地退化是全球干旱和半干旱草地生态系统的共同特征,由于土壤种子库在植被恢复和生物多样性维持等方面的重要作用,已成为研究热点。作为重要的脆弱生态系统,松嫩碱化草甸在人类活动和全球变化影响下发生了明显退化。松嫩碱化草甸土壤种子库研究始于20世纪90年代,主要集中在对虎尾草群落、星星草群落、碱蓬群落、羊草群落等不同演替阶段土壤种子库物种种类、密度、季节变化研究;该地区土壤种子库的物种数目随着恢复演替由初级向高级阶段而呈现逐渐增加的趋势,但物种数目均相对较少,且多以1年生为主。另外对种子雨和某些物种种子散布动态也有零星报道,主要是从群落水平进行的研究,种子雨的空间异质性表现在其组成和大小因群落而异。今后需要加强对松嫩碱化草甸土壤种子库格局与关键生境要素的耦合特征、种子库分布格局的形成机制等研究,研究方法上,引入稳定同位素标记和分子遗传学等研究方法和手段,开展持久种子库的长期定位监测研究,为盐碱地植被恢复提供理论依据。     

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。     

Regeneration of Cyperaceae, with particular reference to seed ecology and seed banks

Cyperaceae (sedges) are an important component of many ecosystems. To understand better their regeneration, we examined seed ecology, including dispersal, seed characteristics, and germination behavior that relate to seed bank development and persistence. We also evaluated sedge seed banks from 104 studies, representing a wide array of habitats. Sedge seed bank development and persistence were associated with germination and dormancy traits, namely: dormancy level, seasonal dormancy patterns, and requirement for light, alternating temperatures, and aerobic conditions. Interplay of traits appears to have resulted in low-risk germination strategies adapted to exploit infrequent occurrence of gaps and facilitate formation of persistent seed banks. A variety of dispersal modes and morphological adaptations occurred, but many species had no apparent specialized structures. The main dispersal vectors were water and then animals. About 216 species, in 21 genera, were recorded in the seed bank survey. High densities (>50,000 m⁻²) occurred occasionally in wet habitats, but generally values were low (<500 m⁻² in 70% of entries). Species richness was also generally low (mean 4.8 species study⁻¹), but ranged from 10 to 33 species in certain wetlands. Our studies showed varied reproductive strategies within habitats, persistence, and ability of many species to colonize disturbed habitats. Overall, seed banks tended to be persistent (>1 year). Maximum longevity ranged between 10 and 295 years, but for certain species viability was lost in <3 years. Seeds of many sedges occurred in deeper soil layers to depths >1 m. Seed production, low in rhizomatous species, ranged between 0 and 345,000 seeds m⁻² year⁻¹. Amphicarpy or pseudo-viviparous plantlets occurred in limited numbers of species. The relation between seed production, seed rain, and seed bank is largely obscure and awaits further investigation. For successful restoration and species conservation projects, seed banks (or a source of seeds) are necessary, combined with suitable germination and establishment conditions. Future seed bank studies are considered.     

Investigations into seed dormancy in Grevillea linearifolia, G. buxifolia and G. sericea: anatomy and histochemistry of the seed coat

BACKGROUND AND AIMS: Seeds of east Australian Grevillea species generally recruit post-fire; previous work showed that the seed coat was the controller of dormancy in Grevillea linearifolia. Former studies on seed development in Grevillea have concentrated on embryology, with little information that would allow testing of hypotheses about the breaking of dormancy by fire-related cues. Our aim was to investigate structural and chemical characteristics of the seed coat that may be related to dormancy for three Grevillea species. METHODS: Seeds of Grevillea linearifolia, Grevillea buxifolia and Grevillea sericea were investigated using gross dissection, thin sectioning and histochemical staining. Water movement across the seed coat was tested for by determining the water content of embryos from imbibed and dry seeds of G. sericea. Penetration of intact seeds by Lucifer Yellow was used to test for internal barriers to diffusion of high-molecular-weight compounds. KEY RESULTS: Two integuments were present in the seed coat: an outer testa, with exo-, meso- and endotestal (palisade) layers, and an inner tegmen of unlignified sclerenchyma. A hypostase at the chalazal end was a region of structural difference in the seed coat, and differed slightly among the three species. An internal cuticle was found on each side of the sclerenchyma layer. The embryos of imbibed seeds had a water content six times that of dry seeds. Barriers to diffusion of Lucifer Yellow existed at the exotestal and the endotestal/hypostase layers. CONCLUSIONS: Several potential mechanisms of seed coat dormancy were identified. The embryo appeared to be completely surrounded by outer and inner barriers to diffusion of high-molecular-weight compounds. Phenolic compounds present in the exotesta could interfere with gas exchange. The sclerenchyma layer, together with strengthening in the endotestal and exotestal cells, could act as a mechanical constraint.     

Seed development and germination ecophysiology of the invasive tree <Emphasis Type="Italic">Prunus serotina</Emphasis> (Rosaceae) in a temperate forest in Western Europe

Seed development, dormancy and germination of the American invasive tree species, Prunus serotina, are described for plants growing in a large forest in Belgium. Seeds of P. serotina were collected following anthesis in the first week of July and thereafter at fortnightly intervals. Seed dormancy, temperature requirements for germination and the soil seed bank were investigated. At maturation (about 105 days after anthesis), seed moisture content had decreased to around 13.7%, and 44% of the seeds had attained the capacity to germinate. Mature seeds of P. serotina exhibited physiological dormancy, germinating only after a long cold, moist stratification period. Highest germination percentage occurred in seeds treated with gibberellic acid (GA₃), at 10°C. We found no evidence that P. serotina forms a persistent seed bank but noticed a persistent seedling bank in the field.     

Seed dormancy release and germination characteristics of <i>Corispermum lehmannianum</i> Bunge,an endemic species in the Gurbantunggut desert of China

Seed dormancy release and germination of Corispermum lehmannianum Bunge were tested using various treatments: temperature, cold stratification, gibberelins (GA3), dry storage and sand burial. Results showed that temperature and light did not affect the germination of fresh seeds, cold stratification and GA3 could improve seed germination, whereas dry storage and sand burial did not. The germination percentage was highest at 35/20 °C after the cold stratification and GA3 treatments. Corispermum lehmannianum seeds were classified as non-deep, Type-2, physiological dormancy (PD), whose seed dormancy could be released by cold stratification and GA3.     

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。     

Seeds photoblastism and its relationship with some plant traits in 136 cacti taxa

Seed germination triggered by light exposure (positive photoblastism) has been determined in quantitative studies for numerous plant families and species. For Cactaceae, positive photoblastism is thought to be associated with life form and seed mass, but this association has never been evaluated. To explore hypotheses on associations between seed mass, seed dispersal, seed dormancy, life form, taxa and plant height with Relative Light Germination (RLG) in Cactaceae, we evaluated the effect of light on seed germination of 136 taxa. The taxa studied are native to several countries: México, Chile, Argentina, Brazil, Perú, USA, and Venezuela. Seed traits contrasted with RLG were life form, seed mass, seed dispersal, seed dormancy, adult plant height and taxon. We found some differences between RLG among taxa; Cacteae, Pachycereeae and Trichocereeae had higher RLG than Notocacteae. RLG was lower for seeds from taller than for shorter taxa, and lower for taxa with heavier seeds than for taxa with lighter seeds. Dispersal syndrome groups varied with RLG. RLG did not differ between cylindrical and globose taxa. Trends found here were in agreement with expectations for small-seeded species to have a light requirement to germinate more often than large-seeded species. This is the first time that cactus height is related to photoblastism. It is possible that seeds from tall plants are larger and thus have the capacity to produce taller seedlings than those from small plants, and that seedlings from large seeds with more resources have the ability to emerge from greater soil depths than those from small seeds.