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Mangroves are one of the most important ecosystems in the world being found in the tropical–subtropical belt. Despite their significance, they have been highly disturbed due to many anthropogenic and natural causes. A significant effort has been made to restore mangroves around the world. However, a lack of information on the seed biology of mangrove species has impeded restoration. Thus, this study aimed to produce a seed dormancy profile for selected plant species of mangroves in Sri Lanka. This profile would allow restoration ecologists to better understand what kinds of dormancy are present, how to alleviate dormancy and how to best stimulate germination to generate seedlings for nursery stock or out-planting. Mature fruits/seeds were collected from coastal zone mangroves in Sri Lanka. Germination and imbibition of non-scarified and manually scarified seeds and embryo:seed length (E:S) ratio of fresh and radicle-emerged seeds were evaluated to assess the class of dormancy. Of the 30 species, seeds from 12 (40%) were non-dormant and 18 (60%) were dormant. Three dormancy classes [physiological (PD), physical (PY) and morphophysiological (MPD)] and presence of epicotyl dormancy were identified. Among species producing dormant seeds, most of them showed PD (44%). PY, MPD and presence of epicotyl dormancy were represented by 28, 17 and 11% of the species, respectively. These findings aid practitioners to craft strategies to effectively break dormancy and germinate seeds for conservation and restoration activities of mangroves.
The objective of this study was to determine how the current (10–16 weeks) and predicted future (2–8 weeks) length of cold stratification and current and predicted future post-stratification temperatures influence radicle and epicotyl emergence in acorns of Quercus robur. We tested radicle and epicotyl emergence at two temperatures corresponding to the current (15/6°C) and predicted future early autumn and spring temperatures (25/15°C) in Poland. We fitted models to describe and derive parameters for radicle and epicotyl emergences over time. The parameters included maximum percentage, rate of emergences, time to achieve the maximum emergence rate, emergence delay and time to 50% emergence. In most cases, the Gompertz model was the best fit, but in a few cases, the logistic model was the best. Richard's model for most of the cases did not converge. This model, according to both information criteria values, was the best fit for epicotyl emergence at 15/6°C following 8 weeks of cold stratification. Richard's model was also the best fit for epicotyl emergence at 25/15°C following 14 weeks of stratification.. Our results indicate that at temperatures typical for early autumn (15/6°C), the time necessary for radicle emergence from 50% of acorns was longer than that from acorns placed at 25/15°C. Four weeks of cold stratification extended 50% radicle emergence at 15/6°C to 70 d, whereas 12 weeks of stratification shortened the time to 11 d. When the acorns were incubated at 25/15°C, radicle emergence occurred faster than at 15/6°C and the time lag between radicle and epicotyl was shorter.
The flora of Mediterranean ecosystems contains families with species having fully and under-developed embryos in their seeds. After-ripening for physiological dormancy release and smoke influence germination in many species. We investigated how after-ripening and embryo growth interact with smoke to influence the temporal dynamics of seedling emergence among fire ephemerals. Seeds were placed in the field and under standardized (50% relative humidity, 30°C) laboratory conditions to test the effects of summer conditions on physiological dormancy loss. Germination was tested with water or smoke compounds (smoke water, KAR1) at a simulated autumn/winter temperature (18/7°C). The timing and amount of seedling emergence with smoke was observed for seeds exposed to near-natural conditions. During summer, physiological dormancy was broken in all species, enabling germination at autumn/winter but not summer temperatures; no embryo growth occurred in seeds with under-developed embryos. At the start of the wet season, seedling emergence from seeds with fully developed embryos occurred earlier than from seeds with under-developed embryos. In a non-consistent manner among our study species, smoke and smoke compounds influenced the rate of embryo growth and amount of germination. Effects of smoke were noticeable in terms of number of emergents in the first emergence season. Among ecologically similar species, we have shown (1) that both thermal and embryo traits exclude germination in the summer, (2) how embryo size influences the timing of seedling emergence in autumn–winter, and (3) a reduced requirement for smoke in the second emergence season after a fire with a shift to reliance on seasonal cues for emergence.
Mangroves are highly adapted to extreme environmental conditions that occur at the interface of salt and fresh water. Adaptations to the saline environment during germination are a key to mangrove survival, and thereby, its distribution. The main objective of this research was to study the effect of salinity on seed germination of selected mangrove species and the application of a hydrotime model to explain the relationship between water potential of the medium and rate of seed germination. Germination of seeds was examined at 15, 25 and 35°C in light/dark over a NaCl gradient. Germination time courses were prepared, and germination data were used to investigate whether these species behave according to the principles of the hydrotime model. The model was fitted for the germination of Acanthus ilicifolius seeds at 25°C. Final germination percentage was significantly influenced by species, osmotic potential and their interaction at 25°C. Moreover, temperature had a clear effect on seed germination (Sonneratia caseolaris and Pemphis acidula) which interacted with osmotic potential. Only A. ilicifolius seeds behaved according to the hydrotime principles and thus its threshold water potential was –1.8 MPa. Optimum germination rates for seeds of the other species occurred at osmotic potentials other than 0 MPa. The descending order of salinity tolerance of the tested species was Aegiceras corniculatum > Sonneratia caseolaris > Acanthus ilicifolius > Pemphis acidula > Allophylus cobbe, suggesting that the viviparous species (A. corniculatum) is highly salt tolerant compared with the non-viviparous species. The results revealed that seeds of the study species exhibited facultative halophytic behaviour in which they can germinate over a broad range of saline environments. Use of a hydrotime model for mangroves was limited as germination of their seeds did not meet model criteria.
Morphophysiological dormancy (MPD) is predominantly found in seeds of temperate regions and is uncommon in arid biomes. MPD has been reported in a number of Hibbertia (Dilleniaceae) species of temperate Australia, and in a single species of the arid zone, H. glaberrima. This study aimed to examine the dormancy and germination ecology of seeds of H. glaberrima. Seeds were subjected to temperature stratification treatments designed to mimic summer and autumn conditions in the Pilbara region of Western Australia. Seed germination and embryo growth were measured. We also tested the interaction between temperature stratification and cycles of drying and wetting designed to mimic sporadic rainfall events. All temperature and moisture treatments were tested in combination (+/–) with the smoke-derived chemical karrikinolide (KAR1). Exposing dormant seeds to temperatures suitable for warm stratification (35°C) for ≥ 8 weeks, followed by incubation at 25°C, resulted in significantly higher germination compared with non-stratified seeds. Exposing seeds to dry/wet cycling in conjunction with temperature stratification did not significantly increase germination. Exposure to KAR1 increased germination under most conditions. Once seeds are shed during October to December, they are exposed to hot and sporadically wet conditions over summer, allowing MPD to be overcome in a proportion of the seed population. Seeds may germinate in autumn (March to April), in conjunction with cooler temperatures. More deeply dormant individuals may require more than one summer to overcome dormancy. Similar to other species occurring in fire-prone ecosystems, fire also plays a crucial role in the germination ecology of H. glaberrima.
Convallariamajalis has double dormancy and hypogeal germination, but no information is available on embryo growth or on the effects of light and gibberellic acid (GA3) on germination in this genus. Therefore, we investigated embryo growth and other germination features in seeds of C. keiskei and compared the data with those of Trillium camschatcense in another study. Until now, in seeds with double dormancy, embryo growth and germination (epigeal) have been studied in detail only for seeds of T. camschatcense. Phenology of embryo growth and emergence of cotyledonary petiole/root (hereafter root) and shoot in seeds of C. keiskei were monitored outdoors. Effects of temperature, light and GA3 on embryo growth and root and shoot emergence were tested under laboratory conditions. Roots emerged the first spring following seed dispersal in autumn. The embryo grew soon after root emergence, and germination was hypogeal. Seeds with an emerged root formed buds from which a shoot (leaf) emerged above ground during the second spring. Alternating temperatures and light had negative effects on root emergence, and GA3 did not substitute for cold stratification in root emergence. Seeds of C. keiskei have double dormancy, but it differs from that in T. camschatcense. Based on differences in embryo growth before (T. camschatcense) versus after (C. keiskei) root emergence, and on epigeal (T. camschatcense) versus hypogeal (C. keiskei) germination, we suggest that two types of deep simple double morphophysiological dormancy (MPD) be recognized. Since embryo growth in C. keiskei does not fit the standard definition of MPD, we propose to expand this definition.
Seeds with epicotyl dormancy reside in soil up to 15 months (or longer), being exposed to a sequence of temperatures, before seedlings completely emerge (i.e. with both roots and shoots). Heretofore, few studies have examined precise temperatures, especially in sequences, for promotion of radicle and cotyledon emergence and how they relate to environmental cues in nature. Viburnum is the best known genus to exhibit epicotyl dormancy and, as such, we investigated the Japanese V.furcatum, hypothesizing a similar kind and level of dormancy. The underdeveloped embryos in mature seeds in October were spatulate shaped, unlike those in other Viburnum species, and they elongated from late June to August of the following year. Radicles emerged after embryo growth until mid-October, followed by cotyledons from mid-April to mid-May. Temperatures required for embryo growth, radicle and cotyledon emergence in the laboratory approximated closely those in the field. Embryo elongation and radicle emergence occurred at warm temperature regimes, and gibberellic acid (GA3) did not substitute for this warm temperature requirements. Following a 120-d cold stratification of seeds with an emerged radicle, shoots emerged from seeds at 10, 15, 15/5, 20/10 and 25/15°C. We identified that seeds of V. furcatum have deep simple epicotyl morphophysiological dormancy like the majority of other Viburnum species. For propagation of the species from seeds, the nearly 2-year period for seedling emergence could be shortened to 8 months: start fresh seeds at 25/15°C (60 d) and then move them through a sequence of 15/5°C (30 d) → 0°C (120 d) → 20/10°C (30 d).
Many temperate plant genera, like Sambucus, have species with range disjunctions among North America, Europe and/or Asia. Cold stratification (sometimes in combination with warm stratification) is the primary mechanism to break seed dormancy in these species. For some of these genera showing Northern Hemispheric disjunctions, members also occur in subtropical or tropical regions, mostly confined to higher elevations where climate and vegetation differ from those in northern latitudes. We made two hypotheses concerning germination for the subtropical Taiwanese Sambucus chinensis: (1) seeds from populations exposed to warm temperatures would require warm stratification, and (2) seeds from populations exposed to cold temperatures need cold stratification. We investigated the germination (including embryo growth) of non-stratified seeds over a range of temperatures and tested the effects of cold stratification and of gibberellins GA3 and GA4 on germination. The amount and timing of germination among populations varied substantially in response to temperature treatments. Seeds from all populations of this species required warm temperatures for dormancy break and germination, regardless of environmental conditions. As such, the majority of seeds had non-deep simple morphophysiological dormancy, which, until now, has not been reported in any members of Sambucus. The seed characteristics of the subtropical S. chinensis are different from those of temperate members of the genus in which cold stratification is the predominate treatment to overcome dormancy.
The purpose of the present study was to investigate the dormancy break and germination requirements of seeds from the rhamnaceous vine Berchemia scandens. The fleshy fruit contains a two-locular stone with an endocarp described as ‘bony, thickish’. Scarified and non-scarified stones increased by about 30–50% in mass during imbibition over a 24-h period. The endocarp of the stone does not completely enclose the seeds and a soft tissue region is present. This region is the primary area of water entrance to the seed, as shown by dye-tracking and by sealing it. Freshly matured and overwintered seeds of B. scandens germinated to low percentages at all temperatures during 2 weeks of incubation in light, and they germinated from moderate to high percentages during 12–14 weeks of incubation in light. While cold stratification had a relatively modest effect on the promotion of total germination across most temperatures assessed (if seeds were left for long enough), it had a somewhat stronger effect on germination rate. Cold-stratified seeds germinated equally well in light and darkness. The class of dormancy found in seeds of B. scandens would be physiological. The anatomy of the stones readily allows water imbibition, showing that seeds of B. scandens lack physical dormancy, an uncommon trait in Rhamnaceae.
Seeds of the narrow-endemic Solidago shortii and of the geographically-widespread S. altissima and S. nemoralis collected and buried in 1992 and 1993 were incubated in light and in darkness at 15/6°, 20/10°, 25/15°, 30/15° or 35/20°C following various periods of burial in soil in a non-temperature-controlled glasshouse. At maturity in November, seeds of the three species germinated to 0–1% in light at 15/6°C and to 10–77% at 20/10°, 25/15°, 30/15° and 35/20°C. Seeds exhumed each April from 1993 to 1996 and incubated in light at 15/6° and 20/10°C germinated to ≥83% and ≥90%, respectively, whereas those exhumed each September of 1993–96 germinated to ≤2%and ≤40%, respectively. At 25/15°, 30/15° and 35/20°C in light, seeds of S. altissima and S. shortii germinated to ≥52% and those of S. nemoralis to ≥19/, regardless of when they were exhumed. Timson's index, which integrates percentages, rates and times for onset of germination, was higher at all temperature regimes for seeds exhumed in April 1995 than for those exhumed in September 1995. Freshly-matured seeds of the three species germinated to 0–11% in darkness. Furthermore, regardless of when they were exhumed, seeds of S. altissima and S. nemoralis incubated in darkness germinated mostly to only 0–9% over the range of temperature regimes. In contrast, ≤88% and ≤6% of the seeds of S. shortii exhumed and incubated in darkness each April and September of 1993–96, respectively, germinated, ≤1% of them germinating while buried in soil. Thus, although buried seeds of all three species exhibited an annual conditional dormancy/non-dormancy cycle, only those of S. shortii exhibited cyclic changes in their germination response in darkness.
The effect of dry storage under ambient laboratory conditions on after-ripening and survivorship was tested on seeds of the geographically-widespread Solidago altissima and S. nemoralis and the narrow-endemic S. shortii. Freshly-matured seeds of S. altissima collected in 1991 and in 1992 germinated to low or moderate percentages in light at 15/6, 20/10 and 25/15°C and to high percentages at 30/15 and 35/20°C, whereas those of S. nemoralis and S. shortii germinated to low percentages over the range of temperature regimes. After 0.8–1.8 years of storage, 1991 seeds of S. altissima incubated in light germinated to high percentages at 25/15, 30/15 and 35/20°C, those of S. nemoralis did so at 30/15 and 35/20°C and those of S. shortii at 20/10, 25/15, 30/15 and 35/20°C; 1992 seeds of all three species germinated to high percentages at 20/10, 25/15, 30/15 and 35/20°C. Freshly-matured 1991 and 1992 seeds of the three species germinated to low percentages in darkness over the range of temperature regimes, and only seeds of S. shortii germinated to high percentages after 0.8–1.8 years of storage. Compared with cold stratification, dry storage was only moderately effective in breaking dormancy in these three species. The primary difference in after-ripening of seeds of the three species was that seeds of the narrow endemic germinated to higher percentages in darkness than those of its two geographically-widespread congeners. Survivorship curves for 1991 and 1992 seeds of S. altissima and S. nemoralis and for 1992 seeds of S. shortii were of Deevey Type I; the survivorship curve for 1991 seeds of S. shortii was closest to Type II. Longevity of 1991 seeds of S. altissima, 1992 seeds of S. nemoralis and 1991 and 1992 seeds of S. shortii was <4.0 years, whereas that of 1991 seeds of S. nemoralis was <2.3 years; 5% of 1992 seeds of S. altissima were viable after 4.0 years.
Seeds of the geographically-widespread Solidago altissima and S. nemoralis and the narrow-endemic S. shortii were buried in pots of soil and placed in a glasshouse without temperature control. After 0.3–4.3 years of burial, some seeds (21–60%) of all three species were viable and they germinated to 75–100% during 2 weeks of incubation in light at 30/15°C. Soil samples collected from several population sites of S. altissima, S. nemoralis and S. shortii were placed in the glasshouse and monitored for seedling emergence. During the first, second, third, fourth, fifth and sixth springs, the number of S. altissima seedlings m−2 emerging was 108–1080, 8–494, 0–520, 0–69, 0–6 and 3, respectively, of S. nemoralis was 108–1122, 17–667, 0–42, 0–6, 0, 0 and 0, respectively, and of S. shortii was 61–1753, 0–25, 0–6, 0, 0 and 0, respectively. More seedlings emerged from disturbed than from non-disturbed soil, but the differences were not significant. Thus, although some seeds of all three species buried in pots remained viable in soil throughout the 4.3-year burial period, longevity was greater and size of seed bank larger in field-collected soil samples containing seeds of the geographically-widespread species than in those containing seeds of the narrow endemic. Although 34 studies have reported seeds of 17 species of Solidago present in soil seed banks, the present study is the first to show, conclusively that Solidago can form a persistent seed bank.
Regardless of whether or not seeds of the geographically-widespread Solidago altissima and S. nemoralis were exposed to light in autumn, those ‘dispersed’ in autumn (15/°C) or winter (5°C) required 12 weeks of light in winter to germinate to ≥80% in darkness in spring (2 weeks at 20/10°C). On the other hand, seeds of the narrow-endemic S. shortii dispersed in autumn and exposed to ≥2 weeks of light in early winter germinated to ≥77% in darkness in spring, and those dispersed in winter and exposed to ≥6 weeks of light germinated to ≥82%. S. altissima and S. nemoralis seeds not exposed to light during any season germinated to only 0–1% in darkness in spring, whereas S. shortii seeds germinated to 45–56%. Seeds of S. altissima and S. nemoralis kept in darkness in autumn and winter needed a 1-day (14-h photoperiod) light exposure in spring to germinate to ≥75% in darkness, whereas those of S. shortii required only one 5-s exposure. Cold-stratified (nondormant) seeds of S. altissima, S. nemoralis and S. shortii exposed to light with a high far-red/red ratio germinated to significantly higher percentages than dark controls and freshly-matured and lab-stored seeds. Results of this study suggest that a soil seed bank of S. shortii should be smaller and be depleted at a faster rate than those of S. altissima and S. nemoralis, and portions of the seeds of the three species can germinate in the far-red-enriched light under plant canopies.
Solidago shortii is endemic to a small area in northcentral Kentucky (USA), whereas two of its sympatric congeners, S. altissima and S. nemoralis, are geographically widespread. Seeds (achenes) of S. shortii (0.370 mg) are significantly larger (PLSD, P=0.05) than those of S. altissima (0.070 mg) and S. nemoralis (0.068 mg). Germination percentages of freshly-matured seeds of the three Solidago species collected in November 1991, 1992 and 1994 were 0–2% in light at 15/6°C, 1–37% at 20/10°C, 9–56% at 25/15°C and 10–85% at 30/15 and 35/20°C. Stratification increased the percentage and rate of germination and decreased the time to the onset of germination (measured by Timson's index only at 20/10°C in light) in the three species. Following 12 weeks of cold stratification in light, seeds of the three species germinated to 72–100% in the light and to 22–100% in darkness over the range of thermoperiods; those cold-stratified in darkness germinated to 39–100% in light. Freshly-matured seeds of S. altissima and of S. nemoralis germinated to 0–4% in darkness, whereas those cold-stratified for 12 weeks in darkness germinated to 0–28% in darkness. On the other hand, freshly-matured and cold-stratified (in darkness) seeds of S. shortii germinated to 0–13 and 13–73%, respectively, in darkness. Under near-natural temperatures in a glasshouse without temperature control, germination of the three species peaked in March. Thus, the primary difference in dormancy-breaking and germination requirements of the three species is that the endemic germinates to a much higher percentage in darkness than its two congeners. Seeds of S. shortii do not have any special dormancy-breaking or germination requirements that could not be fulfilled outside its present-day geographic range.
Seeds of the narrow-endemic Solidago shortii and of the geographically-widespread S. altissima and S. nemoralis buried in December 1993 were exhumed in June 1995 and given 10 cycles of 1 day wet/5 days dry, 9 of 2/5, 8 of 3/5, 7 of 4/5 and 6 of 5/5 in light at 30/15°C; the control was kept continuously wet during the experiment. Seeds of the three species incubated on wet substrate for 3, 4 or 5 days germinated to ≥47% during the first cycle. On the other hand, seeds kept moist 2 days germinated to only 4–26% in the first cycle, and none kept moist for 1 day germinated. Cumulative germination percentages of seeds of all three species at the end of the final cycle of the 1/5 treatment were 0–4%. In the 2/5 treatment, cumulative germination percentages of S. altissima and S. shortii seeds at the end of the final cycle were 50 and 41%, respectively, but that of S. nemoralis was only 4%. For all three species, cumulative germination percentages were ≥55% at the end of the final cycle of the 3/5, 4/5 and 5/5 treatments. Control seeds of the three species germinated to 85–99% after 2 weeks, and no additional seeds germinated during the remainder of the experiment. High percentages of seeds were viable in the treatments and control at the end of the experiment; however, some seeds of S. nemoralis and S. shortii given 1/5, 2/5 and 3/5 treatments became dormant. The ecological implication of this study is that seeds of the three species will not germinate on the soil surface after brief rainfall events in summer. The germination response of the narrow endemic is similar to that of its two geographically-widespread congeners.
Although the underdeveloped embryo, and thus morphological (MD) or morphophysiological (MPD) seed dormancy, is basal in angiosperms, it also occurs in advanced groups. A synthesis of the literature, combining phylogeny and the kind of seed dormancy in the highly evolutionarily advanced order Dipsacales, shows that MPD (or MD) occurs throughout all clades except the most advanced one, Valerina. Seeds of taxa in the Valerina clade have fully developed embryos and physiological dormancy (PD) or are non-dormant (ND); thus, PD and ND are derived conditions in Dipsacales. Assuming that types of seed dormancy have not changed since the Early Tertiary, the fossil record suggests that MPD (or MD) was present in extant genera of Dipsacales by the Palaeocene, but PD (or ND) not until the Miocene. Molecular dating indicates that the ages of dipsacalean lineages with MPD and PD are older than those indicated by the fossil evidence.
Requirements for dormancy break and embryo growth were determined for seeds of the western North American species, Osmorhiza depauperata. Seeds were collected in August 2001 from Sandia Crest (3200 m elevation) and Las Huertas (2300 m), New Mexico (USA). Embryos in fresh seeds were c. 0.6 mm long, and they had to grow to c. 9–10 mm before the radicle emerged from the mericarp. Embryo growth occurred at low temperatures (1 and 5°C), and seeds germinated to high percentages at 1°C during 32 weeks of incubation in the light. No seeds germinated at 5, 15/6, 20/10, 25/15 or 30/15°C during 32 weeks of incubation. Although a 4–18 week warm-temperature (25/15°C) pretreatment increased germination rates at 1°C, it was unnecessary for a high percentage of seeds to germinate. Gibberellic acid (GA3, 10–1000 mg l–1) did not substitute for cold stratification. Seeds from the low-elevation population contained larger embryos and required less time to germinate than those from the high-elevation population. O. depauperata seeds have deep complex morphophysiological dormancy (MPD), which is similar to two other western North American congeners and an Asian congener, but different from two eastern North American congeners. Results from this study suggest that: (1) phylogenetic niche conservatism has played a role in the persistence of deep complex MPD in the three western North American species of Osmorhiza; and (2) the stimulatory effect from a warm pretreatment in species needing only cold stratification for dormancy break is a preadaptation that initiated the development of an absolute warm requirement in species needing both warm and cold stratification.
The most often used time-line for distinguishing a transient seed bank from a persistent seed bank is one calendar year. Thus, species whose seeds live in or on the soil for <1 year have a transient seed bank, whereas those whose seeds live for ≥1 year have a persistent seed bank. However, dormancy cycling of seeds buried in soil has not been given due consideration in these models. When dormancy cycling is considered, it is shown that seeds of both autumn-germinators and spring-germinators are in the dormant state when they are 1 year old. Thus, unless the seeds live until at least the second germination season (i.e. usually 16–18 months following dispersal), they are, in effect, part of a transient seed bank, having lived through only one germination season. We propose that for seeds of such species to be considered part of a short-term persistent seed bank, they should remain viable and germinable until at least the second germination season, and to be part of a long-term persistent seed bank, until at least the sixth germination season. Our definitions are applicable to seeds with physiological, physical or morphophysiological dormancy, which often require >1 year after maturity to come out of dormancy in nature. We discuss modifications of the seedling emergence method for detection of a soil seed bank, so that they correspond to our definitions of seed-bank strategies.