Seed dormancy and germination of the rare,high elevation Balkan endemic <Emphasis Type="Italic">Cerastium dinaricum</Emphasis> (Caryophyllaceae)

Short or long-term ex situ conservation is becoming increasingly important in conservation of plants in today’s changing environments. One of the important steps in ex situ conservation is the collection and storage of seeds and the consequent establishment of seed germination protocols. Cerastium dinaricum (Caryophyllaceae) is an endemic, high elevation and rare species of European conservation concern. Because of its severely fragmented distribution along the Dinaric Alps, the populations are likely to undergo further shrinkage in the future, which addresses the need of a long-term effective conservation management. From the potential ex situ population management perspective, we focused our study on germination ecology of C. dinaricum. The study revealed that temperature considerably affected the germination of seeds, which germinate better at 20 °C rather than 10 °C. A period of cold-wet stratification also significantly improved the final germination percentage with more pronounced increase at 20 °C, while addition of GA3 increased the final germination percentage by breaking the dormancy of non-stratified seeds. Mechanical scarification did not improve germination; on the contrary, it resulted in the lowest germination success. Seeds grown in complete darkness germinated significantly better compared to control when they were exposed to cold-wet stratification. Contrary to previous studies on some alpine species, which germinate better when exposed to light, dark treatment resulted in the highest germination percentages with 70 and 90% germination success after 4 and 8 weeks of stratification, respectively.     

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.     

冷层积和室温干燥贮藏对河西走廊8种荒漠植物种子萌发的影响

不同贮藏和处理条件对不同植物的种子萌发有不同的影响。该文以河西走廊干旱半干旱区8种荒漠植物为研究对象, 探讨了种子经历不同冷层积(4 ℃、-5 ℃、-26 - 10 ℃)和室温干燥贮藏后的萌发响应。研究结果表明: 1)冷层积可使种子萌发率提高、保持不变或降低, 冷层积的有效温度下界可降至-5 ℃或更低。4 ℃和-5 ℃的冷层积使多裂骆驼蓬(Peganum multisectum)和驼蹄瓣(Zygophyllum fabago)种子的萌发率升高、萌发速度加快, 冬季过低的气温以及较大的温度变幅(-26 - 10 ℃)使部分种子萌发率升高。3种冷层积和室温干燥贮藏使黑果枸杞(Lycium ruthenicum)种子萌发率达到90%-100%。唐古特白刺(Nitraria tangutorum)、甘草(Glycyrrhiza uralensis)、苦马豆(Sphaerophysa salsula)种子经过3种冷层积和室温干燥贮藏后萌发率变化较小。中亚紫菀木(Asterothamnus centrali-asiaticus)种子对各种贮藏条件的响应不明显, 部分种子活性丢失。刺沙蓬(Salsola ruthenica)种子扩散时有较高的萌发率(84%), 经-5 ℃和-26 - 10 ℃冷贮藏后, 种子仍具有较高的萌发率, 经4 ℃冷贮藏后几乎不萌发, 大部分种子活性丢失。2)不同物种的种子经过不同方式的贮藏后, 萌发对温度的响应不同。经冷层积后的多裂骆驼蓬种子萌发响应于恒温, 驼蹄瓣和刺沙蓬种子萌发更加响应于变温条件; 多数植物种子在变温培养下萌发速度慢于恒温下。     

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.     

Effect of temperature, light and salinity on seed germination and radicle growth of the geographically widespread halophyte shrub Halocnemum strobilaceum

BACKGROUND AND AIMS: The small leafy succulent shrub Halocnemum strobilaceum occurs in saline habitats from northern Africa and Mediterranean Europe to western Asia, and it is a dominant species in salt deserts such as those of north-west China. The effects of temperature, light/darkness and NaCl salinity were tested on seed germination, and the effects of salinity were tested on seed germination recovery, radicle growth and radicle elongation recovery, using seeds from north-west China; the results were compared with those previously reported on this species from 'salt steppes' in the Mediterranean region of Spain. METHODS: Seed germination was tested over a range of temperatures in light and in darkness and over a range of salinities at 25 degrees C in the light. Seeds that did not germinate in the NaCl solutions were tested for germination in deionized water. Seeds from which radicles had barely emerged in deionized water were transferred to NaCl solutions for 10 d and then back to deionized water for 10 d to test for radicle growth and recovery. KEY RESULTS: Seeds germinated to higher percentages in light than in darkness and at high than at low temperatures. Germination percentages decreased with an increase in salinity from 0.1 to 0.75 M NaCl. Seeds that did not germinate in NaCl solutions did so after transfer to deionized water. Radicle elongation was increased by low salinity, and then it decreased with an increase in salinity, being completely inhibited by > or = 2.0 M NaCl. Elongation of radicles from salt solutions < 3.0 M resumed after seedlings were transferred to deionized water. CONCLUSIONS: The seed and early seedling growth stages of the life cycle of H. strobilaceum are very salt tolerant, and their physiological responses differ somewhat between the Mediterranean 'salt steppe' of Spain and the inland cold salt desert of north-west China.     

The temperature dimension of the seed germination niche in fen wetlands

The seed germination niche partly determines adaptation, ecological breadth and geographic range in plant species. In temperate wetlands, environmental temperature is the chief regulator of germination timing, but the ecological significance of high and low temperatures during dormancy break and germination is still poorly understood. Our aim was to characterize the temperature dimension of the germination niche in mountain base-rich fens, determining (1) the effect of different temperatures on dormancy break and germination, and (2) whether different germination strategies may be identified at the species level. We conducted laboratory germination experiments with seeds of 15 species from these habitats, collected in 18 fen sites in the Cantabrian Mountains (Spain) for two consecutive years. In all the species, the seeds were totally or conditionally dormant at dispersal and stratification produced a significant increase of germination. In most cases, there was not an obligatory requirement for cold temperatures during dormancy break, since warm stratification promoted germination as well. Although the optimal germination thermoperiod was generally high (30/20 °C), most species could also germinate at lower temperatures after cold-stratification. We also identified a group of species associated to cold-water springs that germinated only at low temperatures. Our results demonstrate that dormancy break in mountain base-rich fens does not obligatorily depend on cold temperatures during overwintering. Furthermore, germination at cool temperatures may be more widespread in wetland habitats than previously thought. The existence of two distinctive germination strategies, ‘warm’ and ‘cool’, can potentially give rise to divergent species responses to climate change.     

Variability in seed germination at the interpopulation,intrapopulation and intraindividual levels of the shrub Erica australis in response to fire-related cues

In Mediterranean ecosystems, as well as in other fire prone ones, seedling establishment by some species is particularly favorable right after fire. It has been well established that many plants from these ecosystems have enhanced germination after exposing their seeds to heat or to chemicals related to the passage of fire. Less known it is how variable is this germination response among seeds from different sources (populations, individuals within a population, or parts of the plant within an individual) and whether such differences persist after exposing the seeds to germination-triggering, fire-related cues. In this work we studied the germination response to several fire-related factors of the Mediterranean, lignotuberous shrub Erica australis. This species produces a very combustible fuel, and resprouts vigorously after fire, but conditions for seedling establishment are poorly known. The objective of this study was to evaluate how variable was the germination response of this species among seeds from different sources, and whether this variability changed after exposing the seeds to fire-related factors. Seeds from three different sites, from different individuals at each site, and from different branches within each individual were set to germinate under control conditions or after exposing them to heat or to additions of nitrogenous compounds. Germination was highly variable among populations, individuals within populations or branches within individuals. Exposure to moderate heat promoted germination, but high temperatures were lethal. Differences at the various provenance levels were, however, also observed after heat exposure. Germination of seeds exposed to nitrogenous compounds increased, but differences among populations were always significant. These results indicate that the germination of Erica australis may be linked to disturbances, but it does not seem to be particularly related to passage of fire. On the contrary, this type of response appears more common in plants from heterogeneously disturbed environments. This revised version was published online in August 2006 with corrections to the Cover Date.     

The effect of cold stratification and light on the seed germination of temperate sedges (Carex) from various habitats and implications for regenerative strategies

The germination responses of 32 temperate Carex species were tested in light and darkness at five constant temperatures and under one fluctuating temperature regime, before and after cold-wet stratification. Using a linear logistic regression model, the probability of germination tested across all species was found to be significantly higher after stratification, in light and at the fluctuating temperature. In addition, the probability increased with temperature. Stratification increased germination in 28 species and had very little or no effect on four species. There was almost no germination in darkness prior to stratification, and the germination in light was considerably higher in all but two species compared with that in darkness. Thus, it can be concluded that the Carex species tested have broadly similar germination response patterns. The fact that Carices can be released from high levels of primary dormancy by low-temperature stratification implies that they are spring germinators. A light requirement after stratification in the major fraction of seeds and the capability of almost all investigated sedges to respond to fluctuating temperatures make it likely that persistent seed banks are formed. Additionally, sedges generally seem to have a high temperature requirement for germination which prevents them from emerging at the very beginning of the growing season. Regeneration by seed is probably largely restricted to gaps resulting from late spring disturbances where buried seeds have an opportunity to germinate and grow. Differences in germination were apparent between species occupying different habitats. Overall germination was significantly higher in wetland species than in dry-site species, probably owing to the greater capability of wetland species to respond to fluctuating temperatures. Differences in germination between forest and open-site species can be attributed to the higher capability of forest sedges to respond to low temperatures and temperature fluctuations. The influence of seed weight on germination was not significant in the 18 species adapted to wet, open habitats. There was, however, a tendency for the germination percentages to be low for large-seeded Carices. The interpretation of habitat differences is difficult due to a positive correlation between seed weight and dry habitats.     

Comparisons of germination traits of alpine plants between fellfield and snowbed habitats

We examined the seed-germination responses of 27 alpine species with reference to habitat type (fellfield and snowbed), temperature (five regimes), and light requirement. About 70% of species showed >40% germination at warm temperatures without cold stratification. However, a moist-chilling treatment markedly improved the germination percentages in most species, especially under cool conditions. Thus, cold stratification effectively reduced the temperature requirement for germination. Patterns of germination response within species were not consistent between the fellfield and snowbed habitats for species inhabiting both habitats. For interspecific comparisons, there were no significant differences in germination responses to the temperature regimes and the cold stratification between the fellfield and snowbed species. Also, germination speed and the length of germinating period did not differ between fellfield and snowbed species. Most species (86%) showed a requirement for light for germination without cold stratification. Although the extent of the light requirement was reduced after cold stratification in some species, the light requirement of most small-seeded species remained. The combination of cold stratification and the light requirement is a major factor determining the seedling emergence and formation of seed banks in alpine plants. However, habitat-specific patterns of germination traits were less clear, suggesting similar germination traits in fellfield and snowbed plants, at least under controlled conditions in the laboratory.     

Thermal niche for in situ seed germination by Mediterranean mountain streams: model prediction and validation for Rhamnus persicifolia seeds

Background and Aims

Mediterranean mountain species face exacting ecological conditions of rainy, cold winters and arid, hot summers, which affect seed germination phenology. In this study, a soil heat sum model was used to predict field emergence of Rhamnus persicifolia, an endemic tree species living at the edge of mountain streams of central eastern Sardinia.

Methods

Seeds were incubated in the light at a range of temperatures (10–25 and 25/10 °C) after different periods (up to 3 months) of cold stratification at 5 °C. Base temperatures (T_b), and thermal times for 50 % germination (θ₅₀) were calculated. Seeds were also buried in the soil in two natural populations (Rio Correboi and Rio Olai), both underneath and outside the tree canopy, and exhumed at regular intervals. Soil temperatures were recorded using data loggers and soil heat sum (°Cd) was calculated on the basis of the estimated T_b and soil temperatures.

Key Results

Cold stratification released physiological dormancy (PD), increasing final germination and widening the range of germination temperatures, indicative of a Type 2 non-deep PD. T_b was reduced from 10·5 °C for non-stratified seeds to 2·7 °C for seeds cold stratified for 3 months. The best thermal time model was obtained by fitting probit germination against log °Cd. θ₅₀ was 2·6 log °Cd for untreated seeds and 2·17–2·19 log °Cd for stratified seeds. When θ₅₀ values were integrated with soil heat sum estimates, field emergence was predicted from March to April and confirmed through field observations.

Conclusions

T_b and θ₅₀ values facilitated model development of the thermal niche for in situ germination of R. persicifolia. These experimental approaches may be applied to model the natural regeneration patterns of other species growing on Mediterranean mountain waterways and of physiologically dormant species, with overwintering cold stratification requirement and spring germination.     

Effects of Incubation Temperature on Seed Germination of Mediterranean Plants with Different Geographical Distribution Ranges

Little is known about consequences of the increasing global temperatures on seed germination and even less about the seed germination response of species with different geographical distribution ranges. In the present work, the effect of incubation temperature on seed germination of 31 species from the Iberian Peninsula is analyzed in relation to their geographical distribution range. We tested whether plants with different geographical distribution ranges (Iberian Peninsula endemics, Mediterranean, widely distributed species), respond similarly to incubation temperature using cross-species analysis as well as taking into account phylogenetic relationships among species. Seeds were incubated at 10°C, 15°C, 20°C and 25°C for 6?weeks in darkness. Distribution range appeared as a determinant of germination response to the temperature treatment independent of phylogeny. Widely distributed species were less sensitive than endemic or Mediterranean species to incubation temperatures, which may represent an advantage to colonize new areas or to tolerate changes in the environment. Although their ability to avoid germination during the warm and dry season might be favorable for endemic species in some situations, our results also suggest that under a future of temperature increase imposed by climate change, endemic plant species may be more vulnerable than widely distributed species.     

Seed dormancy in Rumex species in response to environmental factors

Abstract. Many Rumex species show similar seed dormancy characteristics but there is more information concerning R. crispus and R. obtusifolius than other species. These species respond positively to red or white light. Far-red light applied for short periods may promote or inhibit germination depending on the timing of the irradiation in relation to temperature change; but long periods of far-red inhibit germination. Seeds may also be stimulated to germinate in the dark by low-temperature stratification at 15°C or less providing the temperature of the seeds is subsequently raised to a minimum of about 15°C. Seeds can, however, germinate at lower temperatures providing they have received other appropriate stimulatory treatment. Seeds also respond to alternating temperatures. In a diurnal cycle the minimum upper temperature required is about 15°C and the maximum lower temperature is about 25°C. The optimum period spent at the upper temperature is about 8 h when it is 15–25°C but the optimum period decreases as the upper temperature is increased above this range so that at 45°C, for example, it is only about 30 min. The period spent at the lower temperature in a diurnal cycle is not critical. Providing these criteria are met, the percentage germination increases with the number and amplitude of the cycles. The warming part of the cycle is necessary for the response but so far there is no convincing evidence that cooling itself is important. Secondary dormancy is induced at constant temperatures at a rate dependent on temperature, but apparently only in the presence of oxygen. This feature affects the optimum timing of a temperature change or exposure to light. Strong positive interactions are shown between stimulatory temperature treatments and white or red light. Unlike many other weed species the seeds respond only slightly to nitrate ions. The implications of these responses are discussed in relation to field behaviour.     

Seed germination traits of Trapella sinensis (Trapellaceae), an endangered aquatic plant in Japan: Conservation implications

We investigated the effects of cold stratification, temperature, light, and oxygen conditions on seed germination of Trapella sinensis Oliver, an endangered aquatic plant in Japan. Seeds had physiological dormancy, and final germination rate increased with an increasing period of cold stratification. Seeds of T. sinensis had an almost absolute requirement for aerobic conditions to germinate. Also, alternating temperatures significantly promoted germination regardless of light conditions, although final germination percentage was twice to four times higher in light under constant temperature conditions. Suitable sites for germination of T. sinensis appear to be shallow water with alternating temperatures and sufficient oxygen and sediment anoxia, caused by eutrophication of water bodies, may have prevented recruitment of T. sinensis from seed.     

Can seed characteristics or species distribution be used to predict the stratification requirements of herbs in the Australian Alps?

The germination requirements of 19 herbs in the Australian Alps were investigated to determine which species may be sensitive to predicted climate changes. Seeds were subjected to factorial treatments of cold stratification for 0, 4, 8 and 12 weeks, followed by incubation at constant temperatures of 10, 15, 20 and 25 °C and alternating temperatures of 20/5 and 20/10 °C. Germination responses were used to identify stratification‐dependent species, to classify dormancy and to determine optimum conditions for laboratory germination. Ordinal logistic regression was used to determine whether the duration of stratification required for ≥ 50% germination could be predicted by seed weight, seed length, embryo : seed ratio or species distribution (latitudinal range, altitudinal range and maximum altitude). The Kruskal–Wallis test was used to determine any significant differences in stratification requirement between endospermic and non‐endospermic seeds. Species varied considerably in their response to the treatment combinations, and therefore their dormancy class. No significant predictors of stratification requirement were identified by ordinal logistic regression (P > 0.9); however, there was a significant difference in stratification requirement between endospermic and non‐endospermic seeds (P = 0.003). Species with non‐endospermic seeds did not require any stratification to germinate well over a range of temperatures, and appear most likely to remain stable or expand in range in response to climate warming. Conversely, the need for ≥ 8 weeks of cold stratification was associated with the presence of endosperm and either a restricted distribution or upland ecotypes of widely distributed species. Alpine species with endospermic seed and a restricted distribution are most likely to contract in range under climate change and would be appropriate to prioritize for ex situ conservation. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 187–204.     

PHYSIOLOGICAL ECOLOGY OF GERMINATION OF VIOLA RAFINESQUII

Seeds of the winter annual Viola rafinesquii Greene exhibit true dormancy at the time of maturity and dispersal in mid to late spring. During the summer rest period the seeds pass from a state of true dormancy to one of relative dormancy and finally to what may be called a state of complete nondormancy. As the seeds enter relative dormancy they will germinate mostly at relatively low temperatures (10, 15, 15/6, and 20/10 C), but as after-ripening continues they gain the ability also to germinate at higher temperatures (20, 25, and 30/15 C). During June, July, and August seeds will not germinate at field temperatures even if kept continuously moist. But by September and October seeds may germinate to high percentages over a wide range of temperatures, including September and October field temperatures. This pattern of germination responses, involving breaking of true dormancy and widening of the temperature range for germination during relative dormancy, appears to be an adaptation of the species to a hot, dry season. Seeds of V. rafinesquii stored on continuously wet soil (field capacity) or on soil that was alternately wet and dried during the summer did not after-ripen at low temperatures (10, 15, 15/6, and 20/10 C) but did after-ripen fully at high temperatures (20, 25, 30/15, and 35/20 C). Thus, the high temperatures that V. rafinesquii “avoids” by passing the summer in the dormant seed stage actually are required to break seed dormancy and, therefore, are essential for completion of its life cycle.     

Geographic variation in the flood-induced fluctuating temperature requirement for germination in Setaria parviflora seeds

Our aim was to search for specific seed germinative strategies related to flooding escape in Setaria parviflora, a common species across the Americas. For this purpose, we investigated induction after floods, in relation to fluctuating temperature requirements for germination in seeds from mountain, floodplain and successional grasslands. A laboratory experiment was conducted in which seeds were imbibed or immersed in water at 5°C. Seeds were also buried in flood-prone and upland grasslands and exhumed during the flooding season. Additionally, seeds were buried in flooded or drained grassland mesocosms. Germination of exhumed seeds was assayed at 25°C or at 20°C/30°C in the dark or in the presence of red light pulses. After submergence or soil flooding, a high fraction (>32%) of seeds from the floodplain required fluctuating temperatures to germinate. In contrast, seeds from the mountains showed maximum differences in germination between fluctuating and constant temperature treatment only after imbibition (35%) or in non-flooded soil conditions (40%). The fluctuating temperature requirement was not clearly related to the foregoing conditions in the successional grassland seeds. Maximum germination could also be attained with red light pulses to seeds from mountain and successional grasslands. Results show that the fluctuating temperature requirement might help floodplain seeds to germinate after floods, indicating a unique feature of the dormancy of S. parviflora seeds from floodplains, which suggests an adaptive advantage aimed at postponing emergence during inundation periods. In contrast, the fluctuating temperature required for germination among seeds from mountain and successional grasslands show its importance for gap detection.     

A threatened alpine species,Fritillaria tubiformis subsp. moggridgei: Seed morphology and temperature regulation of embryo growth

Abstract

Alpine plants have evolved to fit their life cycle into the short vegetative season of mountain habitats. Fritillaria tubiformis Gren. & Godr. subsp. moggridgei (Boiss. & Reuter ex Planch.) Rix (Liliaceae) is an endemic alpine geophyte, bearing seeds with underdeveloped embryos. Seeds are dispersed in August and embryos complete their development by spring when seeds germinate. In order to optimize seed banking procedures and to develop a proper germination protocol for plant regeneration, we studied embryo morphogenesis and analyzed how this process is influenced by temperature. Radicle protrusion occurred after an incubation of 5 months at 4°C. Under these conditions, underdeveloped embryos reached maturity and acquired a well-defined shoot apex. At the time of dispersal, abundant storage compounds were present in seeds. Lipids and lipid/proteins were uniformly distributed within the embryo and the endosperm, respectively. At late stages of embryo development, starch granules were localized at the cotyledonary tip and were also detected around the shoot meristem. Results suggested that F. tubiformis embryos resumed growth over a large range of temperatures, but were only able to complete development at low temperatures after which they were able to germinate by spring.     

The impact of temperature regimes on development,dormancy breaking and germination of dwarf shrub seeds from arctic,alpine and boreal sites

It has been suggested that the infrequent sexual reproduction of arctic dwarf shrubs might be related to the harsh environmental conditions in which they live. If this is the case, then increases in temperature resulting from global climate change might drastically affect regeneration of arctic species. We examined whether recruitment of Empetrum nigrum ssp. hermaphroditum and Vaccinium uliginosum (hereafter E. nigrum and V. uliginosum) was affected by temperature during three reproductive stages: seed development, dormancy breakage and germination. Seeds were collected from an arctic, an alpine (only E. nigrum) and a boreal site with different climates; stored at different winter temperatures and incubated for germination at different temperatures. Seeds of V. uliginosum developed in the boreal region had a higher percentage germination than did seeds developed in the Arctic. In contrast, seeds of E. nigrum from the arctic site had a higher or similar percentage germination than did seeds from the alpine and boreal sites. Increased winter temperatures had no significant effect on resulting germination percentage of E. nigrum. However, V. uliginosum seeds from the arctic site suffered increased fungal attack (and thus decreased germination) when they were stratified under high winter temperatures. Seeds of both species increased germination with increased incubation temperatures. Our results suggest that both species would increase their germination in response to warmer summers. Longer summers might also favour the slow-germinating E. nigrum. However, increased winter temperatures might increase mortality due to fungal attack in V. uliginosum ecotypes that are not adapted to mild winters.     

Seed dormancy in alpine species

In alpine species the classification of the various mechanisms underlying seed dormancy has been rather questionable and controversial. Thus, we investigated 28 alpine species to evaluate the prevailing types of dormancy. Embryo type and water impermeability of seed coats gave an indication of the potential seed dormancy class. To ascertain the actual dormancy class and level, we performed germination experiments comparing the behavior of seeds without storage, after cold-dry storage, after cold-wet storage, and scarification. We also tested the light requirement for germination in some species. Germination behavior was characterized using the final germination percentage and the mean germination time. Considering the effects of the pretreatments, a refined classification of the prevailing dormancy types was constructed based on the results of our pretreatments. Only two out of the 28 species that we evaluated had predominantly non-dormant seeds. Physiological dormancy was prevalent in 20 species, with deep physiological dormancy being the most abundant, followed by non-deep and intermediate physiological dormancy. Seeds of four species with underdeveloped embryos were assigned to the morphophysiologial dormancy class. An impermeable seed coat was identified in two species, with no additional physiological germination block. We defined these species as having physical dormancy. Light promoted the germination of seeds without storage in all but one species with physiological dormancy. In species with physical dormancy, light responses were of minor importance. We discuss our new classification in the context of former germination studies and draw implications for the timing of germination in the field.     

Drought tolerance and germination response to light and temperature for seeds of saline and non-saline habitats of the habitat-indifferent desert halophyte <Emphasis Type="Italic">Suaeda vermiculata</Emphasis>

Few plants are habitat-indifferent halophytes (i.e., grow well in both saline and non-saline soils). These plants offer a good opportunity to study drought and salinity tolerances during germination for seeds developed and matured in soils differ in salinity. Here, we assessed drought tolerance during germination, as simulated with PEG, and response of germination to light and temperature for Suaeda vermiculata, a habitat-indifferent shrub. Seeds matured in saline and non-saline soils were germinated in six PEG concentrations (0 to ? 1.0 MPa) and put in three incubators adjusted at different temperatures in both light and dark regimes. Drought tolerance was greater for seeds of the non-saline than those of saline soils, especially at higher temperatures. Seeds of the saline soils germinated in the lowest osmotic potentials (? 0.8 and ??1.0 MPa) only at lower temperatures, but seeds of the non-saline soils germinated to higher levels at all temperatures. Tolerances to drought and high temperatures were greater in light for seeds of saline soils, but in darkness for seeds of non-saline soils. Germination rate index did not differ significantly between seeds of the two soil types in higher osmotic potentials, but was significantly greater in seeds of non-saline at lower osmotic potentials. Most seeds that failed to germinate in the PEG concentrations recovered their germination when transferred to distilled water. Germination recovery levels and speeds increased with the decrease in osmotic potentials. Seeds of the saline soil postpone their germination until arrival of suitable temperatures and effective rainfalls that ensure seedling survival in salty habitats of the arid unpredictable deserts.