Dormancy,germination and seed bank storage: a study in support of ex situ conservation of macrophytes of southwest Australian temporary pools

1. Vernal pools and rock pools (gnammas) in the Southwest Australian Floristic Region are forms of temporary wetlands that are under threat. Some of their aquatic macrophytes are rare and/or endemic, and there is a need to develop off‐site seed banks to assure their conservation. Here, we report results of the first comprehensive study of the seed germination, dormancy and seed storage behaviour of nine indigenous macrophyte species. 2. Seeds of Glossostigma drummondii, Myriophyllum balladoniense, M. lapidicola, M. petreaum and Triglochin linearis were non‐dormant, whereas those of Damasonium minus, Glossostigma sp. (currently undescribed), G. trichodes and Myriophyllum crispatum were dormant. Non‐dormant seeds germinated over a range of temperatures (5–20 °C) but temperatures at which highest germination occurred were species specific. All species demonstrated a germination preference for the light. Warm stratification substantially increased germination of dormant Glossostigma trichodes seeds and all dormancy‐breaking treatments partially overcame dormancy in Glossostigma spp. 3. Seeds possessed orthodox storage behaviour (tolerating drying to 5% moisture content and storage at ?18 °C) and are thus amenable to seed banking as a means of ex situ conservation. 4. It appears that seeds of most species are able to germinate upon inundation as long as they are situated at the soil surface. Thus, species are opportunistic and respond to the first rains of the season providing prompt ecological cuing in an environment vulnerable to rapid drying events. Maintaining the integrity of the soil crust may be an important first step for on‐site conservation if seeds are in the superficial layers.     

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.     

Variation within species and inter-species comparison of seed dormancy and germination of four annual Lamium species

In an ecological context, knowledge of intra-species variation in dormancy and germination is necessary both for practical and theoretical reasons. We used four or five seed batches (replicates) of four closely related annuals co-occurring in arable fields in Sweden: Lamium amplexicaule, L. confertum, L. hybridum and L. purpureum. Seeds used for experiments stemmed from plants cultivated on two sites, each site harbouring one population of each species, thereby ensuring similar environmental history of seeds. Seeds were tested for germination when fresh and after three different pre-treatments (cold or warm stratification, or dry storage) for up to 24 weeks. Seeds were also sown outdoors. Despite substantial intra-species variation, there were clear differences between species. The general seed dormancy pattern, i.e. which environmental circumstances that affect dormancy, was similar for all species; dormancy reduction occurred during warm stratification or dry storage. Even though the response to warm stratification indicates a winter annual pattern, successful plants in Sweden were mostly spring emerged. Germination in autumn occurred, but plants survived winters poorly. Consequently, as cold stratification did not reduce dormancy, strong dormancy in combination with dormancy reduction during dry periods might explain spring germination. It is hypothesised that local adaptations occur through changes mainly in dormancy strength, i.e. how much effort is needed to reduce dormancy. Strong dormancy restricts the part of each seed batch that germinate during autumn, and thus reduces the risk of winter mortality, in Sweden.     

Variation in dormancy and germination in three co-occurring perennial forest herbs

Mesic deciduous forest herbs often disperse seed with morphophysiological dormancy (MPD) that prevents germination during unfavorable periods for seedling survival. However, for seeds of some species with MPD, seasonal separation of root and shoot emergence and variation in dormancy levels can complicate interpretation of seedling emergence timing in the field. We tested whether dormancy-break and germination requirements differed among co-occurring perennial forest herbs, Actaea racemosa, Hydrastis canadensis, and Sanguinaria canadensis, which are wild-harvested for their medicinal properties and known to have MPD. Seeds of all species exhibited a summer → autumn → winter requirement for seedling emergence in spring. However, species differed in seed-bank persistence due to variation in primary dormancy levels and stratification requirement of seeds. A. racemosa and H. canadensis can form short-term persistent seed bank, whereas S. canadensis can form a long-term persistent seed-bank, regardless of whether elaiosomes were removed from seeds prior to burial. A. racemosa seeds are dispersed in autumn with weak physiological dormancy, as seeds germinated to high rates at 15/6°C after 8 weeks. In contrast, most seeds of the summer dispersed species, H. canadensis and S. canadensis, require summer temperatures to overcome physiological dormancy. Consequently, seedling emergence is reduced and delayed by 1 year if seeds are not sown immediately following the period of natural dispersal. Seedling emergence was much lower in the field than in controlled conditions for all species, especially in the small-seeded A. racemosa. Interspecific variation in dormancy levels and germination traits must be considered when establishing populations for conservation purposes and in understanding recruitment limitation in perennial forest herbs.     

Seed germination ecology of the threatened endemic Iberian <Emphasis Type="Italic">Delphinium fissum</Emphasis> subsp. <Emphasis Type="Italic">sordidum</Emphasis> (Ranunculaceae)

Seeds of Delphinium fissum subsp. sordidum are physiologically dormant at maturity, with underdeveloped embryos; thus they have morphophysiological dormancy (MPD). The aims of this study were to determine the requirements for embryo growth, dormancy break and germination, to characterise the type of seed dormancy and to evaluate the effects of light, seed age, pollination mechanism, and inter-annual and inter-population variability on germinative ability. After 3 months of incubation at 5°C (cold stratification) in darkness conditions, the mean embryo length increased from 5.6 to 2.07 mm, with 76% of seeds germinating. Conversely, embryos of seeds incubated during 3 months at 20/7 or 28/14°C hardly grew and no germination was recorded. Since cold stratification was the only requirement for the loss of MPD, and both dry storage in laboratory conditions and warm stratification prior to cold stratification shortened the cold stratification period required for germination, it could be concluded that D. fissum subsp. sordidum seeds have intermediate complex MPD. Cold stratification and incubation in darkness conditions promoted higher germination percentages than those in light. In addition, germinative ability increased with seed age up to 8 months (reaching 96% at 5°C in darkness), showed a pronounced inter-annual and inter-population variability, as well as a significant decrease in seeds coming from pollination by geitonogamy. High temperatures (25/10 or 28/14°C) induced seeds to secondary dormancy, so seedling emergence in the greenhouse was restricted to February–March. The requirements for dormancy break and germination reflect an adaptation to trigger germination in late winter. This study is the first one to document a gradual increase in germination percentage with seed age for plant species with intermediate complex MPD.     

Hormonal and environmental regulation of seed germination in flixweed (<Emphasis Type="Italic">Descurainia sophia</Emphasis>)

Flixweed is one of the most abundant weeds in North America and China, and causes a reduction in crop yields. Dormancy of flixweed seeds is deep at maturity and is maintained in soil for several months. To identify regulators of seed dormancy and germination of flixweed, the effect of environmental and hormonal signals were examined using dormant and non-dormant seeds. The level of dormancy was decreased during after-ripening and stratification, but long imbibition (over 5 days) at 4 °C in the dark resulted in the introduction of secondary dormancy. The strict requirement of duration of cold treatment for the break of dormancy may play a role in the seasonal regulation of germination. The germination of non-dormant flixweed seeds was critically regulated by red (R) and far-red (FR) light in a photoreversible manner. Sodium nitroprusside, a donor of nitric oxide (NO), promoted germination of half-dormant seeds, suggesting that NO reduced the level of seed dormancy. As has been shown in other related species, light elevated sensitivity to GA₄ in dark-imbibied flixweed seeds, but cold treatment did not affect GA₄-sensitivity unlike in Arabidopsis. Taken together, our results indicate that seed germination in flixweed and its close relative Arabidopsis is controlled by similar as well as distinct mechanisms in response to various endogenous and environmental signals.     

Dormancy and germination of sixRhinanthus species in relation to climate

The genusRhinanthus (Orobanchaceae) consists of annual hemiparasites that occur in a wide range of climates. Patterns of dormancy and germination were studied for six species sampled in areas ranging from the Pyrenees to Northern Scandinavia, and from sea level up to about 2500 m altitude in the Alpine region. Dormancy was broken by a 2 to 6 months period of cold stratification. Optimal temperature and length of the stratification period appeared to vary between and within species. Two patterns of dormancy and germination were distinguished. Seeds of the first group, including the widely distributedR. minor andR. angustifolius, further referred to as the LW group, require a relatively long period of cold stratification. Moreover, their germination is accelerated if they are subjected to a widening range of higher temperatures in the last weeks of the stratification period. In the other species (R. alectorolophus, R. glacialis, R. mediterraneus and perhapsR. antiquus) the release of dormancy is completed in a rather short period. Higher temperatures in the last weeks of the stratification period hardly affect the germination process of this SN-group, with short dormancy and no accelerated germination at higher temperatures. In both groups, temperatures above the low values during stratification reduce germination percentages and induce secondary dormancy in non-germinated seeds. The effects strongly vary with timing and temperature. The patterns have a genetic basis and seem to be species-specific. They do not vary with climate conditions, since samplesR. minor, whether collected in sub-arctic or sub-alpine areas or at sea level, generally react according to the LW pattern, and samples ofR. alectorolophus from areas at sea level up to montane regions according to the SN one. The ecological significance of the various stratification requirements and effects of higher temperatures on germination is discussed in relation to the local climate of the species and the evolutionary history of the genus.     

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。     

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.     

Achene germination of the spring ephemeroid species Carex physodes in the Gurbantunggut Desert

Much of the seed germination research on Carex has focused on wetland species, and little is known about the species of arid habitats. Here, we investigated seed dormancy and germination of Carex physodes, which is an important component of the plant community of the Gurbantunggut Desert of the Junggar Basin in Xinjiang, China. Our studies included the effects of mechanical and chemical scarification, dry storage, treatment with GA₃, wet‐cold stratification, and burial in the field. No freshly matured achenes germinated over a range of temperature regimes after treatment with GA₃, 6 months of dry storage or removal of part of the endosperm. The mechanical scarification resulted in < 5% achene germination, however, higher percentage of achene germination occurred after removal of the pericarp (60%), H₂SO₄ scarification (30%) or scarification in 10% NaOH (85%). Six and nine months of wet‐cold stratification promoted < 40% achene germination. The optimal germination temperatures ranged from 25/10°C to 35/20°C. Maximum germination after 9 months of burial at a depth of 3 cm in the field was 36%. These results indicate that the seeds have non‐deep physiological dormancy (PD) and that the pericarp contains germination inhibitors and has strong mechanical resistance to germination.     

The comparative germination ecology of nine Rumex species

The germination requirements, dormancy cycle and longevity of nine Rumexspecies were studied in field conditions and laboratory experiments to show theadaptations of the related species to their specific habitat. Within one genus,rather striking differences were observed in germination ecology. However, theclosely related species, R. acetosa and R.scutatus, are very similar: they fruit in early summer; theirseeds can germinate immediately after dispersal, and they are nondormant andshort-lived. R. acetosella also has fruits insummer, but the seeds do not germinate the first season after dispersal. Theyare long-lived, but buried seeds do not show a dormancy cycle; they mightgerminate in different seasons after exposure to light. Seeds of four species (R. conglomeratus,R. maritimus, R. sanguineus andR. crispus) are long-lived and undergo aseasonal dormancy cycle, with a low level of dormancy in winter and early springand a deep dormancy in summer as was already known for R.obtusifolius. These seeds are shed in the autumn, and they germinatemainly in the spring in consecutive years. R. maritimusalso germinates in summer and autumn on drying muddy soils. The seeds of R. hydrolapathum only germinate onwaterlogged soils, which explains its growth at the edge of streams and ponds.Its seeds are rather short-lived. The seeds of the species on very wetplaces require a higher temperature for germination.     

Tools for <Emphasis Type="Italic">Carex</Emphasis> revegetation in freshwater wetlands: understanding dormancy loss and germination temperature requirements

Carex is a globally distributed genus with more than 2000 species worldwide and Carex species are the characteristic vegetation of sedge meadow wetlands. In the mid-continental United States, Carex species are dominant in natural freshwater wetlands yet are slow to recolonize hydrologically restored wetlands. To aid in Carex revegetation efforts, we determined the dormancy breaking and temperature germination requirements of 12 Carex species. Seeds were cold stratified at 5/1°C for 0–6 months and then incubated in light at 5/1°C, 14/1°C, 22/8°C, 27/15°C, or 35/30°C. We found that all Carex species produced conditionally dormant seeds. The optimal temperature for germination for all but three species was 27/15°C. As is the case in other species with physiological dormancy, cold stratification increased germination percentages, broadened the temperature range suitable for germination, and increased germination rates for most species, but the magnitude of the effects varied among species. Many species germinated to 80% at 27/15°C without cold stratification and at 22/8°C with ≤1 month of stratification but required much longer stratification (up to 6 months depending on the species) to germinate to 80% at 14/1°C and 35/30°C. Our findings illustrate how a stratification pretreatment can greatly benefit Carex seed sowing efforts by triggering rapid germination to higher percentages. We recommend that cold stratification be targeted towards species with strong dormancy or used across a wider range of species when seed supplies for restoration are limiting. For Carex revegetation, establishing Carex canopies rapidly may help to prevent the invasion of undesirable species such as Phalaris arundinacea.     

Seed germination requirements of Buck’s beard [Aruncus dioicus (Walter) Fernald (Rosaceae)]

Aruncus dioicus (Walter) Fernald (Rosaceae) is a perennial herbaceous plant whose young shoots are traditionally collected in the wild and consumed as a food in NE Italy. The aim of this study was to determine the germination requirements of its seeds in order to start its cultivation, and to assess the germination of six accessions of the species. Viability of seeds ranged from 86 to 97% in the various accessions. Germination rate was almost null in seeds of two accessions, and ranged from 10.5 to 37.3 in the other ones. The seed coat was permeable to water. Treatments with GA₃, KNO₃ and mechanical scarification did not enhance the germination, while the cold stratification treatment at 2 °C for different periods improved the germination rate and the mean germination time as compared with the untreated seeds. With 45 days of cold stratification, the germination rate and mean germination time (respectively, 90.1% and 7.7 dd) of seeds were different from those of the untreated seeds. Cold stratified seeds germinated under artificial light and did not germinate in the dark. Seeds of A. dioicus displayed an intermediate physiological dormancy, removable by a cold stratification treatment, requiring both light and cold conditions.     

Seed germination patterns in green dragon (Arisaema dracontium,Araceae)

Arisaema dracontium (green dragon) is a perennial herb that is widely distributed in eastern North America. However, in Canada, at the northern edge of its distribution, the species is designated as “vulnerable” with respect to conservation status. In natural populations, seedlings are uncommon; the present study was undertaken in order to characterize seed and seedling properties in green dragon. Seeds were sampled from five sites, ranging from Ontario at the northern limit of the distribution range, to Louisiana in the south. Seed germinability ranged from 25 to 55%, depending upon source. Experiments indicated that neither the hard seed coat nor a water-soluble exudate from the seed was responsible for inducing or maintaining dormancy. Patterns of seed germination appear to reflect general climatic conditions at the sites where seeds had originated. Cold stratification at 3°C produced significantly greater relative germinability in all seed collections except the most southerly one, from Baton Rouge. These seeds also had a slower overall speed of germination. In contrast, germination of seeds from the most northerly site was promoted by cold stratification and occurred over a relatively brief period. Germination in alternating light and dark conditions decreased the speed of germination compared to germination in the dark, however exposure to light changed the phenology of germination by promoting development of adventitious roots and primary leaves in these seedlings.     

Comparative <Emphasis Type="Italic">in vitro</Emphasis> germination ecology of <Emphasis Type="Italic">Calopogon tuberosus</Emphasis> var. <Emphasis Type="Italic">tuberosus</Emphasis> (Orchidaceae) across its geographic range

Seed responses to temperature are often essential to the study of germination ecology, but the ecological role of temperature in orchid seed germination remains uncertain. The response of orchid seeds to cold stratification have been studied, but the exact physiological role remains unclear. No studies exist that compare the effects of either cold stratification or temperature on germination among distant populations of the same species. In two separate experiments, the role of temperature (25, 22/11, 27/15, 29/19, 33/24°C) and chilling at 10°C on in vitro seed germination were investigated using distant populations of Calopogon tuberosus var. tuberosus. Cooler temperatures promoted germination of Michigan seeds; warmer temperatures promoted germination of South Carolina and north central Florida seeds. South Florida seed germination was highest under both warm and cool temperatures. More advanced seedling development generally occurred at higher temperatures with the exception of south Florida seedlings, in which the warmest temperature suppressed development. Fluctuating diurnal temperatures were more beneficial for germination compared to constant temperatures. Cold stratification had a positive effect on germination among all populations, but South Carolina seeds required the longest chilling treatments to obtain maximum germination. Results from the cold stratification experiment indicate that a physiological dormancy is present, but the degree of dormancy varies across the species range. The variable responses among populations may indicate ecotypic differentiation.     

Seed germination behavior of five Resedaceae species from arid Arabian desert: gibberellic acid and light effects

Abstract

Light and growth-promoting compounds, such as gibberellic acids (GA₃), are among the most important factors that can break physiological seed dormancy. Here, we investigate the effects of GA₃ and light on germination of five species of Resedaceae that are known to have different levels of physiological dormancy. Seeds were incubated at 20/30?°C in both 12-hr photoperiod and complete darkness. To study the effect of growth hormone on germination, seeds were soaked for 24?h in different concentrations of GA₃ before sowing. The annuals (Reseda aucheri and Oligomeris linifolia) and the perennial Ochradenus arabicus had deep physiological dormancy, and exogenous application of GA₃ enhanced their germination in the light, but not in darkness; few or no seeds germinated in the dark in these species. Ochradenus aucheri and O. baccatus had intermediate and non-deep dormancy, respectively, and application of GA₃ enhanced their germination in both light and darkness. Germination of the annual species was much slower than that of the shrubby perennials. Overall, these results indicate that conditions under which seed developed, matured and stored on maternal plants as well as incubation conditions should be taken into consideration when assessing germination behavior of the perennial species of Resedaceae.     

Comparative germination ecology of two altitudinal vicariant Saxifraga species endemic to the north of Spain

Seeds of high‐mountain species are thought to germinate rapidly, synchronously and at high percentages after a cold period, with limited dependence on the external environment; yet, empirical evidence only partially supports this behaviour. We performed a comparative study of the germination response of two closely related taxa along an altitude gradient in northern Spain. Seeds from several maternal families of six populations of Saxifraga trifurcata (lowland species) and S. canaliculata (highland species) were subjected to temperature and stratification treatments. Germination percentages and germination rates were analysed using generalised linear mixed modelling and accelerated failure‐time modelling. We found that germination percentages and germination rates were high and dependent on incubation temperature in both species. Within species, seeds from higher altitudes had higher germination percentages under all conditions. Cold–wet stratification negatively affected germination success, particularly in the lowland species. Overall, the highland species was less responsive to the experimental treatments and showed more synchronous germination patterns. We conclude that seeds from these two Saxifraga species germinate as efficiently as species from other habitats, but have a narrower germination response, probably due to the stronger selective pressures in their harsh environments. Finally, a cold, wet stratification period is not a prerequisite for the germination of high‐mountain S. canaliculata, and its strong negative effect on the germination of its lowland relative S. trifurcata may contribute to the altitudinal segregation of these two species.     

Interpopulation variability on embryo growth,seed dormancy break,and germination in the endangered Iberian daffodil Narcissus eugeniae (Amaryllidaceae)

The main goal of the study was to assess germination requirements in a threatened daffodil to elaborate a detailed protocol for plant production from seeds, a key tool for conservation. Experiments were carried out both in the laboratory and outdoor conditions. In Pseudonarcissi section, endemic Iberian species of Narcissus studied heretofore have different levels of morphophysiological dormancy (MPD). Embryo length, radicle emergence, and shoot emergence were analyzed to determine the level of MPD. Both interpopulational variability and seed storage duration were also studied. Mean embryo length in fresh seeds was 1.32 mm and the embryo had to grow until it reached at least 2.00 mm to germinate. Embryo growth occurs during warm stratification, after which the radicle emerges when temperatures go down. Seed dormancy was broken in the laboratory at 28/14°C in darkness followed by 15/4°C, but the germination percentage varies depending on the population. In outdoor conditions, seed dispersal occurs in June, the embryo grows during the summer and then the radicle emerges in autumn. The radicle system continues to grow during the winter months, but the shoot does not emerge until the beginning of the spring because it is physiologically dormant and requires a cold period to break dormancy. Early cold temperatures interrupt embryo growth and induce dormancy in seeds with an advanced embryo development. Seeds of N. eugeniae have deep simple epicotyl MPD. In addition, we found that embryo growth and germination were improved by seed storage duration.     

Seed germination of the carnivorous plant Byblis gigantea (Byblidaceae) is cued by warm stratification and karrikinolide

Byblidaceae is one of the most poorly studied carnivorous plant families, with seed dormancy and germination biology remaining unresolved. This knowledge deficit has significant conservation and management implications, particularly as the most southerly distributed species, the south‐west Western Australian endemic Byblis gigantea, is listed as critically endangered. This study examined the ecophysiology of seed dormancy and germination in B. gigantea in concert with a study of the population dynamics of a single plant community. Mass seedling emergence and plant establishment were observed after a wildfire in 2007 in the natural population, followed by a rapid decline in mature individuals (91%) over a 4‐year monitoring period, with almost no inter‐fire recruitment (1.4% of all emergence) observed. Seeds possessed a fully developed embryo, and the germination characteristics of fresh seeds classified them as showing physiological dormancy. Seed dormancy was partially alleviated by warm stratification (30 °C) for 8 weeks prior to incubation at 15 °C, with c. 40% germination observed. With the additional exposure of seeds to the germination‐active chemical in smoke, karrikinolide, the germination of warm‐stratified seeds increased to 89%. Seeds also displayed orthodox storage behaviour and appeared to be amenable to long‐term seed banking for conservation. These results present the first observation of the stimulation of the germination of a carnivorous plant by a smoke‐derived compound. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 143–152.     

Caryopsis germination and seedling emergence in an inland dune dominant grass Leymus secalinus

Leymus secalinus (Georg.) Tzvel. (Poaceae) is a dominant sand dune grass inhabiting the Mu-Us Sandland, semiarid China. Freshly harvested caryopses (seeds) are in non-deep physiological dormancy (non-deep PD) because of low percentage and slow rate of germination. Experiments were conducted to examine the effects of temperature, cold stratification, caryopsis coat scarification or partial removal of endosperm and sand burial on caryopsis dormancy, germination and seedling emergence. Caryopsis germination was significantly influenced by duration of cold stratification, temperature and their interactions. After 8 weeks of cold stratification, caryopsis germination percentage at 30 °C reached to 90%, equally in light or darkness. Rate and percentages of germination were also hastened and increased by scarifying the caryopsis coat or by artificial removal of different proportions of the endosperm. However, seedling developmental characteristics were significantly influenced by the proportion of the endosperm that remained in the caryopses. Seedling emergence, caryopsis germination and enforced dormancy in sand were significantly affected by sand burial depth. As sand burial depth increased, caryopsis germination and seedling emergence decreased whereas caryopsis enforced dormancy increased. 1–2 cm was the optimal depths for caryopses germination and seedling emergence. Although there were still 30% caryopses germinated at 8 cm, the maximal burial depth for seedling emergence was only 4 cm. The partial germination strategy regulated by non-deep PD, temperature and sand burial ensures that only a few caryopses germinated each time and may reduce risks for seedling survival.