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.     

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.     

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.     

喀什霸王的结实和种子萌发特性

喀什霸王(Zygophyllum kaschgaricum)是生长于中国新疆南部荒漠环境的稀有种及二级保护植物。当前, 该物种在自然种群中呈分散式及片段化分布, 且种群密度低, 种群老龄化较严重。因此, 为了了解该物种种子萌发特性及其对荒漠环境的响应, 该文采用室内控制实验方法, 对该物种的自然坐果率、结籽率、种子吸水特性、种子休眠和萌发特性及对干旱胁迫的响应进行了比较研究。结果表明: 喀什霸王在自然种群中的坐果率及结籽率较低, 且种子的败育率较高。不同干藏时间种子的吸水速率间存在显著差异; 随着干藏时间的延长, 种子的吸水率逐步增强。刚成熟的种子在不同温度及光周期下均可萌发; 其中高温(10/20 ℃, 20/30 ℃)及黑暗条件下的萌发率比低温(10/5 ℃, 5/2 ℃)及光照条件下的萌发率高。不同干藏时间的种子在不同浓度赤霉素(GA₃)下的萌发率均较高; 但低温储藏时间对该物种种子的打破休眠及萌发率没有促进作用。以上结果说明该物种存在非深度生理休眠; 而干藏时间、高温且黑暗及高浓度(50 mmoloL ^-1) GA₃是打破休眠及促进种子萌发的最合适条件。高温条件下的干旱胁迫对喀什霸王种子萌发具有抑制作用; 春季和秋季降水量决定种子的萌发率。总之, 喀什霸王种子在物候上表现出的春秋季萌动及非深度生理休眠以提高幼苗存活力及保障种群稳定性, 是一种对新疆南部干旱及高温胁迫荒漠环境的适应策略。     

The impact of global warming on germination and seedling emergence in Alliaria petiolata,a woodland species with dormancy loss dependent on low temperature

• The impact of global warming on seed dormancy loss and germination was investigated in Alliaria petiolata (garlic mustard), a common woodland/hedgerow plant in Eurasia, considered invasive in North America. Increased temperature may have serious implications, since seeds of this species germinate and emerge at low temperatures early in spring to establish and grow before canopy development of competing species.
• Dormancy was evaluated in seeds buried in field soils. Seedling emergence was also investigated in the field, and in a thermogradient tunnel under global warming scenarios representing predicted UK air temperatures through to 2080.
• Dormancy was simple, and its relief required the accumulation of low temperature chilling time. Under a global warming scenario, dormancy relief and seedling emergence declined and seed mortality increased as soil temperature increased along a thermal gradient. Seedling emergence advanced with soil temperature, peaking 8 days earlier under 2080 conditions.
• The results indicate that as mean temperature increases due to global warming, the chilling requirement for dormancy relief may not be fully satisfied, but seedling emergence will continue from low dormancy seeds in the population. Adaptation resulting from selection of this low dormancy proportion is likely to reduce the overall population chilling requirement. Seedling emergence is also likely to keep pace with the advancement of biological spring, enabling A. petiolata to maintain its strategy of establishment before the woodland canopy closes. However, this potential for adaptation may be countered by increased seed mortality in the seed bank as soils warm.

     

Temperature requirements for dormancy break and seed germination vary greatly among 14 wetland Carex species

We evaluated dormancy loss in seeds of 14 Carex species (C. atherodes, C. brevior, C. comosa, C. cristatella, C. cryptolepis, C. granularis, C. hystericina, C. lacustris, C. pellita, C. scoparia, C. stipata, C. stricta, C. utriculata, C. vulpinoidea) under growing season and stratification conditions and determined the temperature requirements for germination. Seeds were germinated for 1 year at a diel temperature regime (5/1 °C, 14/1 °C, 22/8 °C, or 27/15 °C) or a seasonal regime (seeds moved among the four diel regimes to mimic seasonal temperatures). All species had conditionally dormant seeds at maturity. The optimal temperature for germination of most species was 27/15 °C. The 14 species were grouped by their seed viability, dormancy, and germination with a Seed Regeneration Index (SRI; range 0–1) using the results of this study and a previously published paper on stratification effects on Carex seed dormancy and germination. The eight species that had an SRI value >0.5 (C. brevior, C. comosa, C. cristatella, C. cryptolepis, C. hystericina, C. scoparia, C. stipata, C. vulpinoidea) had high seed viability (>60%) and required little to no stratification to germinate readily over a broad range of temperatures. The six species with an SRI value <0.5 (C. atherodes, C. granularis, C. lacustris, C. pellita, C. stricta, C. utriculata) generally had low seed viability (<50% and often <1%) and required stratification or particular temperatures (35/30 °C or 5/1 °C for C. stricta; 35/30 °C for C. utriculata; 27/15 °C for C. atherodes, C. lacustris, C. pellita; 5/1 °C for C. granularis) for germination ≥50%. These six species will require more attention from restoration practitioners to ensure that there are sufficient viable seeds to meet revegetation goals, that dormancy break is achieved, and that seeds are sown when temperatures are optimal for germination. The different seed germination syndromes that we found for these Carex species likely contribute to variable seed bank formation and emergence patterns, and species coexistence.     

Sympatric species of Hibbertia (Dilleniaceae) vary in dormancy break and germination requirements: implications for classifying morphophysiological dormancy in Mediterranean biomes

Background and Aims

Several ecologically important plant families in Mediterranean biomes have seeds with morphophysiological dormancy (MPD) but have been poorly studied. The aim of this study was to understand the seed ecology of these species by focusing on the prominent, yet intractably dormant Australian genus Hibbertia. It was hypothesized that the slow germination in species of this genus is caused by a requirement for embryo growth inside the seed before germination, and that initiation of embryo growth is reliant upon a complex sequence of environmental cues including seasonal fluctuations in temperature and moisture, and an interplay with light and smoke. Using the results, the classification of the MPD level in species of Hibbertia is considered.

Methods

Four species of Hibbertia in winter rainfall south-western Australia were selected. These species, whilst differing in geographic distributions, are variously sympatric, and all are important understorey components of plant communities. The following aspects related to dormancy break, embryo growth and germination were investigated: temperature and moisture requirements; effects of karrikinolide, gibberellic acid and aerosol smoke; and phenology.

Key Results

Following exposure to wet/dry cycles at low or high temperatures, embryo growth and germination occurred, albeit slowly in all species at low temperatures when moisture was unlimited, corresponding to winter in south-west Australia. Photo regime influenced germination only in H. racemosa. Aerosol smoke triggered substantial germination during the 1st germination season in H. huegelii and H. hypericoides.

Conclusions

Although the study species are con-generic, sympatric and produce seeds of identical morphology, they possessed different dormancy-break and germination requirements. The physiological component of MPD was non-deep in H. racemosa but varied in the other three species where more deeply dormant seeds required >1 summer to overcome dormancy and, thus, germination was spread over time. Embryos grew during winter, but future studies need to resolve the role of cold versus warm stratification by using constant temperature regimes. To include Mediterranean species with MPD, some modifications to the current seed-dormancy classification system may need consideration: (a) wet/dry conditions for warm stratification and (b) a relatively long period for warm stratification. These outcomes have important implications for improving experimental approaches to resolve the effective use of broadcast seed for ecological restoration.     

Epicotyl morphophysiological dormancy in seeds of Lilium polyphyllum (Liliaceae)

Dormancy-breaking and seed germination studies in genus Lilium reveal that the majority of Lilium spp. studied have an underdeveloped embryo at maturity, which grows inside the seed before the radicle emerges. Additionally, the embryo, radicle or cotyledon has a physiological component of dormancy; thus, Lilium seeds have morphophysiological dormancy (MPD). A previous study suggested that seeds of Lilium polyphyllum have MPD but the study did not investigate the development of the embryo, which is one of the main criteria to determine MPD in seeds. To test this hypothesis, we investigated embryo growth and emergence of radicles and epicotyls in seeds over a range of temperatures. At maturity, seeds had underdeveloped embryos which developed fully at warm temperature within 6 weeks. Immediately after embryo growth, radicles also emerged at warm temperatures. However, epicotyls failed to emerge soon after radicle emergence. Epicotyls emerged from >90% seeds with an emerged radicle only after they were subjected to 2 weeks of cold moist stratification. The overall temperature requirements for dormancy-breaking and seed germination indicate a non-deep simple epicotyl MPD in L. polyphyllum.     

Effects of seed maturation time and dry storage on light and temperature requirements during germination in invasive Prosopis juliflora

The effects of time of seed maturation and dry seed storage and of light and temperature requirements during seed incubation on final germination percentage and germination rate were assessed for the invasive shrub Prosopis juliflora (Sw.) D.C., grown under desert environmental conditions of the United Arab Emirates (UAE). Seeds were collected from Fujira on the northern coast of the UAE at different times during the growing seasons (autumn, winter and spring) and were germinated immediately and after 8 months of dry storage under room temperature (20±3 °C). Seeds were germinated at three temperatures (15, 25 and 40 °C) in both continuous light and darkness. The results showed significant effects for time of seed collection, seed storage, light and temperature of seed incubation and many of their interactions on both germination percentage and rate. Fresh seeds matured during autumn and winter germinated significantly greater at 40 °C and in light than at lower temperatures and in dark. Storage significantly increased germination percentage and rate; the increase was greater for seeds matured during winter than for seeds matured during spring. This indicates that dormancy breakage was greater in seeds of winter than seeds of spring. The need for high temperature to achieve greater germination was significantly reduced after seed storage, especially for seeds matured in autumn and winter.     

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.     

Climate warming could shift the timing of seed germination in alpine plants

Background and Aims

Despite the considerable number of studies on the impacts of climate change on alpine plants, there have been few attempts to investigate its effect on regeneration. Recruitment from seeds is a key event in the life-history of plants, affecting their spread and evolution and seasonal changes in climate will inevitably affect recruitment success. Here, an investigation was made of how climate change will affect the timing and the level of germination in eight alpine species of the glacier foreland.

Methods

Using a novel approach which considered the altitudinal variation of temperature as a surrogate for future climate scenarios, seeds were exposed to 12 different cycles of simulated seasonal temperatures in the laboratory, derived from measurements at the soil surface at the study site.

Key Results

Under present climatic conditions, germination occurred in spring, in all but one species, after seeds had experienced autumn and winter seasons. However, autumn warming resulted in a significant increase in germination in all but two species. In contrast, seed germination was less sensitive to changes in spring and/or winter temperatures, which affected only three species.

Conclusions

Climate warming will lead to a shift from spring to autumn emergence but the extent of this change across species will be driven by seed dormancy status. Ungerminated seeds at the end of autumn will be exposed to shorter winter seasons and lower spring temperatures in a future, warmer climate, but these changes will only have a minor impact on germination. The extent to which climate change will be detrimental to regeneration from seed is less likely to be due to a significant negative effect on germination per se, but rather to seedling emergence in seasons that the species are not adapted to experience. Emergence in autumn could have major implications for species currently adapted to emerge in spring.     

Seed dormancy and germination in Jeffersonia dubia (Berberidaceae) as affected by temperature and gibberellic acid

The genus Jeffersonia, which contains only two species, has a trans‐Atlantic disjunct distribution. The aims of this study were to determine the requirements for breaking dormancy and germination of J. dubia seeds and to compare its dormancy characteristics with those of the congener in eastern North America. Ripe seeds of J. dubia contain an underdeveloped embryo and were permeable to water. In nature, seeds were dispersed in May, while embryos began to grow in September, and were fully elongated by late November. Germination started in March of the next year, and seeds emerged as seedlings soon after germination. In laboratory experiments, incubation at high temperatures (25 °C, 25/15 °C) for at least 8 weeks was required to initiate embryo growth, while a transfer to moderate temperatures (20/10 °C, 15/6 °C) was needed for the completion of embryo growth. At least 8 weeks at 5 °C was effective in overcoming physiological dormancy and for germination in seeds after the embryos had fully elongated. Thus, both high and low temperatures were essential to break dormancy. Gibberellic acid (GA₃) treatment could substitute for the high temperature requirement, but not for the low temperature requirement. Based on the dormancy‐breaking requirements, it is confirmed that the seeds have deep simple morphophysiological dormancy. This dormancy type is similar to that of seeds of the eastern North American species J. diphylla. Although seeds require 10–11 months from seed dispersal to germination in nature, under controlled conditions they required only 3 months after treatment with 1000 mg·l^?1 GA₃, followed by incubation at 15/6 °C. This represents practical knowledge for propagation of these plants from seed.     

Effect of Temperatures on Dormancy and Germination in Three Species in the Lamiaceae Occurring in Northern Wetlands

In the temperate region temperature is the main factor influencing the germination period of plant species. The purpose of this study was to examine effects of constant and fluctuating temperatures on dormancy and germination under laboratory and field conditions in the three wetland species Lycopus europaeus, Mentha aquatica and Stachys palustris. The results should give indications if the temperature-dependent regulation of dormancy and germination is phylogenetically constrained. Tests for germination requirements showed a minimum temperature for germination of 9 °C in Mentha and 12 °C in Lycopus and Stachys, and a maximum temperature of 33 °C for Lycopus and 36 °C for Mentha and Stachys. Fluctuating temperatures promoted germination in all three species but the amplitude required for high germination (>50%) differed: it was 8 °C in Mentha, 10 °C in Stachys and 14 °C in Lycopus (mean temperature 22 °C). The effect of temperatures on the level of dormancy was examined in the laboratory by imbibing seeds at temperatures between 3 °C and 18 °C for periods between 2 and 28 weeks, as well as by a 30-month burial period, followed by germination tests at various temperatures, in light and darkness. In the laboratory only low temperatures (≤12 °C) relieved primary dormancy in seeds of Lycopus, while in Mentha and Stachys also higher temperatures lead to an increase of germination. Dormancy was only induced in Lycopus seeds after prolonged imbibition at 12 °C in the laboratory. Buried seeds of all species exhibited annual dormancy cycles with lower germination in summer and higher germination from autumn to spring. Exhumed seeds, however, showed considerable differences in periods of germination success. Dormancy was relieved when ambient temperatures were below 12 °C. Ambient temperatures that caused an induction of dormancy varied depending on species and test condition, but even low temperatures (8 °C) were effective. At high test temperatures (25 °C) in light, exhumed seeds of all three species showed high germination throughout the year. The three species showed various differences in the effects of temperatures on dormancy and germination. Similarities in dormancy and germination found among the species are in common with other spring-germinating species occurring in wetlands, so it seems that the temperature dependent regulation of dormancy and germination are related to habitat and not to phylogenetic relatedness.     

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

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

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.     

Seed Germination Biology of the Narrowly Endemic Species Lesquerella stonensis (Brassicaceae)

Abstract Lesquerella stonensis (Brassicaceae) is an obligate winter annual endemic to a small portion of Rutherford County in the Central Basin of Tennessee, where it grows in disturbed habitats. This species forms a persistent seed bank, and seeds remain viable in the soil for at least 6 years. Seeds are dormant at maturity in May and are dispersed as soon as they ripen. Some of the seeds produced in the current year, as well as some of those in the persistent seed bank, afterripen during late spring and summer; others do not afterripen and thus remain dormant. Seeds require actual or simulated spring/summer temperatures to come out of dormancy. Germination occurs in September and October. Fully afterripened seeds germinate over a wide range of thermoperiods (15/6–35/20°C) and to a much higher percentage in light (14 h photoperiod) than in darkness. The optimum daily thermoperiod for germination was 30/15°C. Nondormant seeds that do not germinate in autumn are induced back into dormancy (secondary dormancy) by low temperatures (e.g., 5°C) during winter, and those that are dormant do not afterripen; thus seeds cannot germinate in spring. These seed dormancy/ germination characteristics of L. stonensis do not differ from those reported for some geographically widespread, weedy species of winter annuals and thus do not help account for the narrow endemism of this species.     

Tanned or Burned: The Role of Fire in Shaping Physical Seed Dormancy

Plant species with physical seed dormancy are common in mediterranean fire-prone ecosystems. Because fire breaks seed dormancy and enhances the recruitment of many species, this trait might be considered adaptive in fire-prone environments. However, to what extent the temperature thresholds that break physical seed dormancy have been shaped by fire (i.e., for post-fire recruitment) or by summer temperatures in the bare soil (i.e., for recruitment in fire-independent gaps) remains unknown. Our hypothesis is that the temperature thresholds that break physical seed dormancy have been shaped by fire and thus we predict higher dormancy lost in response to fire than in response to summer temperatures. We tested this hypothesis in six woody species with physical seed dormancy occurring in fire-prone areas across the Mediterranean Basin. Seeds from different populations of each species were subject to heat treatments simulating fire (i.e., a single high temperature peak of 100°C, 120°C or 150°C for 5 minutes) and heat treatments simulating summer (i.e., temperature fluctuations; 30 daily cycles of 3 hours at 31°C, 4 hours at 43°C, 3 hours at 33°C and 14 hours at 18°C). Fire treatments broke dormancy and stimulated germination in all populations of all species. In contrast, summer treatments had no effect over the seed dormancy for most species and only enhanced the germination in Ulex parviflorus, although less than the fire treatments. Our results suggest that in Mediterranean species with physical dormancy, the temperature thresholds necessary to trigger seed germination are better explained as a response to fire than as a response to summer temperatures. The high level of dormancy release by the heat produced by fire might enforce most recruitment to be capitalized into a single post-fire pulse when the most favorable conditions occur. This supports the important role of fire in shaping seed traits.     

A season- and gap-detection mechanism regulates seed germination of two temperate forest pioneers

The survival of seedlings in temperate climate habitats depends on both temporal and spatial factors. The interaction between an internal seed dormancy mechanism and the ruling environmental conditions allows accurate cueing of germination. We analysed how environmental signals interact in seeds of temperate forest pioneer species, increasing the seed's chances of germinating in the right place at the right time. Digitalis purpurea and Scrophularia nodosa are two small-seeded herbaceous species that typically grow in vegetation gaps in European temperate forests. Seeds of both species are partially dormant at the time of dispersal in summer. This primary dormancy is released in autumn and early winter, resulting in a minimal level of physiological dormancy by late winter and early spring. We observed that physiological dormancy was induced again in seeds exhumed in late spring and in summer. Experiments in laboratory conditions revealed that primary dormancy in seeds of S nodosa was broken by cold stratification, whereas primary dormancy in D. purpurea seeds was broken by both a cold and a warm stratification. The two species differed in their response to the tested gap-detection signals, as light was the most important factor stimulating germination of D. purpurea, and seeds of S. nodosa germinated best when subjected to daily fluctuating temperatures. This study clearly indicates that the ability to germinate in response to gap-detection signals changes seasonally in temperate forest pioneers. Additionally, seeds of both species responded differently to these environmental signals, probably reflecting differences in the regeneration niche.     

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.     

Temperature requirements differ for the two stages of seed dormancy break in Aegopodium podagraria (Apiaceae), a species with deep complex morphophysiological dormancy

Only a few studies have considered the possibility that low temperature requirements may vary among stages of dormancy break in seeds with morphophysiological dormancy (MPD). We show that this lack of consideration in previous studies on seed dormancy and germination of Aegopodium podagraria might explain the low germination percentages and/or the relatively long periods of incubation needed for germination. Under natural temperatures, embryos began to grow in September and were fully elongated by late December; most growth occurred when the average daily mean temperature was about 10°C. Radicles emerged under snow in late winter, and cotyledons emerged after snowmelt in early spring. In laboratory experiments, 100% of the embryos grew to full length at both 0 and 5°C, whereas 0°C was much more effective than 5°C in overcoming the physiological dormancy in seeds after embryos were fully elongated. Following radicle emergence, cotyledons emerged readily in a wide range of temperatures ≥5°C. GA(3) did not substitute for the low temperature requirement for dormancy break. Seed dormancy in A. podagraria fits Nikolaeva's formula for deep complex MPD, i.e., C(3)B-C(3). Better germination of seeds pretreated at 0° than at 5°C has practical implications for cultivating this species.