Temperature effects on embryonic abscisic acid levels during development of wheat grain dormancy

Abscisic acid (ABA) levels were determined in both the embryo and remaining grain remnant during development of wheat caryopses under temperature conditions which produced either high or low levels of dormancy in mature grain. Higher levels of grain dormancy were produced in grain from plants grown at 15°C as compared to 25°C. In grain grown at 15°C, embryonic ABA levels steadily increased during development, reaching a maximum at stage IV, just before grain desiccation. At 25°C, ABA levels were very high at the earliest stages of embryonic development, but dropped rapidly during maturation. Only small cultivar differences in ABA levels were observed during development at either temperature. In general, higher levels of dormancy in mature grain correlated with prolonged elevation of ABA levels during grain maturation.     

Hormonal and temperature regulation of seed dormancy and germination in <Emphasis Type="Italic">Leymus chinensis</Emphasis>

Fluctuating temperature plays a critical role in determining the timing of seed germination in many plant species. However, the physiological and biochemical mechanisms underlying such a response have been paid little attention. The present study investigated the effect of plant growth regulators and cold stratification in regulating Leymus chinensis seed germination and dormancy response to temperature. Results showed that seed germination was less than 2 % at all constant temperatures while fluctuating temperature significantly increased germination percentage. The highest germination was 71 % at 20/30 °C. Removal of the embryo enclosing material of L. chinensis seed germinated to 74 %, and replaced the requirement for fluctuating temperature to germinate, by increasing embryo growth potential. Applications of GA₄₊₇ significantly increased seed germination at constant temperature. Also, inhibition of GA biosynthesis significantly decreased seed germination at fluctuating temperatures depending upon paclobutrazol concentration. This implied GA was necessary for non-dormant seed germination and played an important role in regulating seed germination response to temperature. Inhibition of ABA biosynthesis during imbibition completely released seed dormancy at 20/30 °C, but showed no effect on seed germination at constant temperature, suggesting ABA biosynthesis was important for seed dormancy maintenance but may not involve in seed germination response to temperature. Cold stratification with water or GA₃ induced seed into secondary dormancy, but this effect was reversed by exogenous FL, suggesting ABA biosynthesis during cold stratification was involved in secondary dormancy. Also, cold stratification with FL entirely replaced the requirement of fluctuating temperature for germination with seeds having 73 % germination at constant temperature. This appears to be attributed to inhibition of ABA biosynthesis and an increase of GA biosynthesis during cold stratification, leading to an increased embryo growth potential. We suggest that fluctuating temperature promotes seed germination by increasing embryo growth potential, mainly attributed to GA biosynthesis during imbibitions. ABA is important for seed dormancy maintenance and induction but showed less effect on non-dormant seed germination response to temperature.     

Hypoxia interferes with ABA metabolism and increases ABA sensitivity in embryos of dormant barley grains

Two mechanisms have been suggested as being responsible for dormancy in barley grain: (i) ABA in the embryo, and (ii) limitation of oxygen supply to the embryo by oxygen fixation as a result of the oxidation of phenolic compounds in the glumellae. The aim of the present work was to investigate whether hypoxia imposed by the glumellae interferes with ABA metabolism in the embryo, thus resulting in dormancy. In dormant and non-dormant grains incubated at 20 degrees C and in non-dormant grains incubated at 30 degrees C (i.e. when dormancy is not expressed), ABA content in the embryo decreased dramatically during the first 5 h of incubation before germination was detected. By contrast, germination of dormant grains was less than 2% within 48 h at 30 degrees C and embryo ABA content increased during the first hours of incubation and then remained 2-4 times higher than in embryos from grains in which dormancy was not expressed. Removal of the glumellae allowed germination of dormant grains at 30 degrees C and the embryos did not display the initial increase in ABA content. Incubation of de-hulled grains under 5% oxygen to mimic the effect of glumellae, restored the initial increase ABA in content and completely inhibited germination. Incubation of embryos isolated from dormant grains, in the presence of a wide range of ABA concentrations and under various oxygen tensions, revealed that hypoxia increased embryo sensitivity to ABA by 2-fold. This effect was more pronounced at 30 degrees C than at 20 degrees C. Furthermore, when embryos from dormant grains were incubated at 30 degrees C in the presence of 10 microM ABA, their endogenous ABA content remained constant after 48 h of incubation under air, while it increased dramatically in embryos incubated under hypoxia, indicating that the apparent increase in embryo ABA responsiveness induced by hypoxia was, in part, mediated by an inability of the embryo to inactivate ABA. Taken together these results suggest that hypoxia, either imposed artificially or by the glumellae, increases embryo sensitivity to ABA and interferes with ABA metabolism.     

Conditions of induction of secondary dormancy in apple after breaking of primary dormancy by anaerobiosis and low temperature

Dormant embryos of Pyrus malus L. cv. Golden delicious, isolated from the fruits at harvest time or after a few months storage at 10 to 15°C, were kept under anaerobic conditions in order to eliminate primary dormancy. Germination tests were then carried out at different temperatures, using three modes of culture depending on the nature of the contact between the embryo and the medium. In CM the distal part of the two cotyledons was immersed in the medium. In RM only the embryonic axis was immersed. In C/2M the embryo was placed flat on the medium, the radicle and the external surface of one cotyledon being in contact with it.
Results showed that primary dormancy was released progressively depending on the duration of the anaerobic treatment. After a treatment of 11 or 13 days the last symptoms of primary dormancy were only apparent when germination tests were carried out at high temperatures (26–30°C) or in CM mode of culture.
When the embryos were kept at 4°C during 3 months inside the fruits, subsequent germination was inhibited at high temperature and in CM mode of culture. When the embryos were kept under anaerobic conditions (7 days) after the chilling treatmem inside the fruits, germination was no longer inhibited. It is concluded that the inhibition of germination at high temperature and in CM mode of culture is due to the persistence of traces of primary dormancy. Therefore, these conditions do not seem to induce secondary dormancy in apple embryos.
After elimination of primary dormancy by anaerobiosis. only application of (±) abscisic acid (3.8 and 19 μM) inhibited germination. These results support the idea that ABA is an important factor in the induction of dormancy. However, the question remains whether this secondary embryo dormancy has the same characteristics as the original primary dormancy.     

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.     

Seed dormancy in Acer: Is there a common mechanism for all Acer species and what part is played in it by abscisic acid?

Seeds of Acer pseudoplatanus L. are usually considered to show only testa-imposed dormancy, but a transient embryo dormancy has also been identified at the time of fruit dispersal. Even embryos that did not show full dormancy at this stage possessed low germinative vigour. Removal of embryo dormancy and the development of increased germination potential did not require the low temperatures necessary for the removal of testa-imposed dormancy in this species. Germination rates of embryos from freshly harvested or briefly stored fruits were accelerated by removal of cotyledonary tissue, the most rapid responses occurring in isolated embryonic axes following complete removal of both cotyledons. Longer storage reduced this effect because of the increases in germinative vigour of whole embryos. Abscisic acid (ABA) reinforced embryo dormancy in embryos from freshly harvested or briefly stored fruits and also reduced germination rates in similarly derived isolated embryonic axes. This response to ABA also became progressively less marked as the storage period was extended. Loss of embryo dormancy was correlated with a reduction in endogenous abscisic acid levels in both whole embryos and cotyledons, suggesting that endogenous ABA contributes to the regulation of embryo dormancy in these seeds. There are no indications, however, that endogenous ABA is directly implicated in the low temperature processes associated with the removal of testa-imposed dormancy. The relevance of embryo dormancy in the intact seed of A. pseudoplatanus is discussed.     

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 abscisic acid and stratification on somatic embryo maturation and germination of holm oak (<Emphasis Type="Italic">Quercus ilex</Emphasis> L.)

Summary The effect of abscisic acid (ABA) was evaluated during the maturation and germination of holm oak (Quercus ilex L.) somatic embryos. The addition of ABA to the culture medium significantly reduced unwanted recurrent embryogenesis in mature somatic embryos without affecting the germination of embryos subjected to stratification at 4°C. Stratification at 4°C for 2 mo. was the most efficient for stimulating somatic embryo germination of holm oak. The addition of 90 and 450 mM sucrose also improved germination, while higher sucrose concentrations were inhibitory.     

Temperature effects on embryonic abscisic acid levels during development of wheat grain dormancy

Abscisic acid (ABA) levels were determined in both the embryo and remaining grain remnant during development of wheat caryopses under temperature conditions which produced either high or low levels of dormancy in mature grain. Higher levels of grain dormancy were produced in grain from plants grown at 15°C as compared to 25°C. In grain grown at 15°C, embryonic ABA levels steadily increased during development, reaching a maximum at stage IV, just before grain desiccation. At 25°C, ABA levels were very high at the earliest stages of embryonic development, but dropped rapidly during maturation. Only small cultivar differences in ABA levels were observed during development at either temperature. In general, higher levels of dormancy in mature grain correlated with prolonged elevation of ABA levels during grain maturation.Contribution from USDA-ARS and the College of Agriculture and Home Economics Research Center, Washington State University, scientific paper no. 8901-13.Mention of a specific product name by the United States Department of Agriculture does not constitute an endorsement and does not imply a recommendation over other suitable products.     

Effects of Stratification, Gibberellic acid and Germination Temperature on the Germination of Betula nana

Experiments were carried out with three seed lots of Betula nana collected in 1967 from different localities in Norway. Seeds were stratified for 0-20 days in dark at +2-+3 °C on filter papers moistened with distilled water, or treated with solution of GA₃ for 24 h at room temperature, and then moved into special germination boxes that were placed in different temperature conditions. All the seed lots had conditional dormancy. Quantitatively, the dormancy was different in the different seed lots (pronenances), but there were no qualitative difference in the reaction to stratification gibberellic acid and to germination temperature. Differences between seed lots may have been due to different stage of seed development. The dormancy was deepest at low temperatures(12 and 15°C) decreasing gradually with increasing temperature (to 24 °C). The dormancy was effectively broken by a short stratification (from 5 to 15 days), and by treatment with gibberellic acid. The deeper the dormancy and the lower the germination temperature the longer the stratification that was needed for maximum germination. Similarly, the concentration of GA₃ needed for maximum germination increased with decreasing temperature and with increasing dormancy.     

Involvement of ABA in induction of secondary dormancy in barley (Hordeum vulgare L.) seeds

At harvest, barley seeds are dormant because their germination is difficult above 20 degrees C. Incubation of primary dormant seeds at 30 degrees C, a temperature at which they do not germinate, results in a loss of their ability to germinate at 20 degrees C. This phenomenon which corresponds to an induction of a secondary dormancy is already observed after a pre-treatment at 30 degrees C as short as 4-6 h, and is optimal after 24-48 h. It is associated with maintenance of a high level of embryo ABA content during seed incubation at 30 degrees C, and after seed transfer at 20 degrees C, while ABA content decreases rapidly in embryos of primary dormant seeds placed directly at 20 degrees C. Induction of secondary dormancy also results in an increase in embryo responsiveness to ABA at 20 degrees C. Application of ABA during seed treatment at 30 degrees C has no significant additive effect on the further germination at 20 degrees C. In contrast, incubation of primary dormant seeds at 20 degrees C for 48 and 72 h in the presence of ABA inhibits further germination on water similarly to 24-48 h incubation at 30 degrees C. However fluridone, an inhibitor of ABA synthesis, applied during incubation of the grains at 30 degrees C has only a slight effect on ABA content and secondary dormancy. Expression of genes involved in ABA metabolism (HvABA8'OH-1, HvNCED1 and HvNCED2) was studied in relation to the expression of primary and secondary dormancies. The results presented suggest a specific role for HvNCED1 and HvNCED2 in regulation of ABA synthesis in secondary seed dormancy.     

Accumulation and leakage of abscisic acid during embryo development and seed dormancy in wheat

Seed dormancy develops latein embryogenesis after a period of potential prematuregermination and has been associated with levels ofabscisic acid (ABA) in, and sensitivity to, ABA ofembryos. In wheat (Triticum aestivum L.)embryos, there are two peaks in levels of ABA duringdevelopment: the first occurs 25 days afterpollination (DAP) and the second from 35 to 40 DAP. The first peak of ABA appears to be associated withthe development of the embryo's sensitivity to ABAsince such sensitivity was altered in seeds on earsthat were incubated in a solution of ABA from 15 and20 DAP. In the embryos of Kitakei wheat, a line thatexhibits dormancy, the second peak, at around 35 DAP,was more prolonged in comparison to Chihoku, anon-dormant line. The results support the proposedinvolvement of ABA in the formation and maintenance ofseed dormancy during middle and late embryogenesis. When developing embryos were incubated in water,embryonic ABA leaked out from the embryos, inparticular between 30 and 40 DAP. Prematuregermination observed between 30 and 40 DAP might berelated to such leakage of ABA from embryos.     

Changes in Abscisic Acid Biosynthesis and Catabolism during Dormancy Breaking in Fagus sylvatica Embryo

At harvest, embryos of Fagus sylvatica are dormant. A cold pretreatment without medium at 30% moisture content allowed them to germinate. A comparison of the abscisic acid (ABA) content before and after the pretreatment has no significant relevance since dormancy is expressed during the culture at 23°C. During this culture, both de novo biosynthesis and conjugate hydrolysis contributed to maintain a high level of ABA in the dormant axis. The level of conjugates and the rate of hydrolysis were not modified substantially by the cold pretreatment. In contrast, the dormancy release was associated with a strong decrease in the capacity for ABA synthesis. Moreover, feeding (+)-[³H]ABA to untreated and pretreated embryos proved that the cold treatment also induced a hastening of ABA catabolism. Received August 15, 1996; accepted December 6, 1996     

Predicting patterns of post-fire germination in 35 eastern Australian Fabaceae

Germination in 35 species from 15 legume genera of southeastern Australia was promoted by a heat treatment which broke the seed coatcaused dormancy. Once the critical temperature was reached, most seeds had their dormancy broken, independent of the duration of heating. Species fell into three classes according to whether their dormancy was broken by a temperature of 40, 60 or 80°C. Highest germination in all species was achieved by heating in the temperature range 80–100°C, although long durations (120 min) at 100°C caused seed death in several species. At 120°C, seeds of most species were killed at all but one minute's duration. A proportion of seeds from 7 species (Acacia myrtifolia, Pultenaea daphnoides, P. incurvata, P. linophylla, P. polifolia, Dillwynia floribunda and Sphaerolobium vimineurn) was not killed at 120°C and had their dormancy broken. This proportion varied markedly and resultant germination levels were significantly less than those at 80 and 100°C, except in S. vimineum. Between-site variations in the 4 species tested (A. myrtifolia, A. suaveolens, A. terminalis and A. ulicifolia) were small. These variations concerned: (i) the minimum temperature required to break seed dormancy in 2 species: 60°C in one population of A. myrtifolia and A. suaveolens, and 80°C in the other; and (ii) the intensity of the germination response. Duration of heating was less important than temperature as a determinant of germination. Ordination techniques revealed that results from one duration across temperatures were comparable with data from multiple durations. This has significant applications in studying rare species, where seed may be in short supply. Predicted germination levels after a moderate intensity fire should far exceed those after a low intensity fire. Little germination was predicted for many species after a low intensity fire and for one species, A. elongata, no germination was predicted. The potential role of indicator species in relation to the maintenance of species in a community is suggested.     

Response of Amaranthus retroflexus L. seeds to gibberellic acid, ethylene and abscisic acid depending on duration of stratification and burial

Freshly harvested, dormant seeds of Amaranthus retroflexus were unable to germinate at 25 and 35 °C. To release their dormancy at the above temperatures, the seeds were stratified at a constant temperature (4 °C) under laboratory conditions or at fluctuating temperatures in soil or by outdoor burial in soil. Fully dormant, or seeds stratified or buried (2006/2007 and 2007/2008) for various periods were treated with exogenous gibberellic acid (GA₃), ethephon and abscisic acid (ABA). Likewise, the effects of these regulators, applied during stratification, on seed germination were determined. The results indicate that A. retroflexus seed dormancy can be released either by stratification or by autumn–winter burial. The effect of GA₃ and ethylene, liberated from ethephon, applied after various periods of stratification or during stratification, depends on dormancy level. GA₃ did not affect or only slightly stimulated the germination of non-stratified, fully dormant seeds at 25 and 35 °C respectively. Ethylene increased germination at both temperatures. Seed response to GA₃ and ethylene at 25 °C was increased when dormancy was partially removed by stratification at constant or fluctuating temperatures or autumn–winter burial. The response to GA₃ and ethylene increased with increasing time of stratification. The presence of GA₃ and ethephon during stratification may stimulate germination at 35 °C. Thus, both GA₃ and ethylene can partially substitute the requirement for stratification or autumn–winter burial. Both hormones may also stimulate germination of secondary dormant seeds, exhumed in September. The response to ABA decreased in parallel with an increasing time of stratification and burial up to May 2007 or March 2008. Endogenous GA_n, ethylene and ABA may be involved in the control of dormancy state and germination of A. retroflexus. It is possible that releasing dormancy by stratification or partial burial is associated with changes in ABA/GA and ethylene balance and/or sensitivity to these hormones.     

Peroxidases in Relation to Removal of Dormancy and Germination of Apple Embryos

The changes in germination, peroxidase activity and isoperoxidase spectrum have been studied in apple embryos at 5°C (stratification) and at 20°C in the presence or absence of seed coats. The embryo dormancy is progressively released at 5°C, but not at 20°C. The peroxidase activity in embryos covered with seed coats is very low at 5°C as well as at 20°C which corresponds to a restricted number of isoenzymes. In isolated embryos the peroxidase activity increases significantly. This is due to an increase in both the number and the activity of the isoperoxidases and it is more pronounced at 20°C than at 5°C. The obtained results suggest that the soluble peroxidases are not involved in the process of the release of embryo dormancy. The variations observed are attributed to the growth process following germination, which can occur even at low temperature.     

Effect of grain colour gene (R) on grain dormancy and sensitivity of the embryo to abscisic acid (ABA) in wheat

The level of grain dormancy and sensitivity to ABA of the embryo, a key factor in grain dormancy, were examined in developing grains of a white-grained wheat line, Novosibirskaya 67 (NS-67), and its red-grained near-isogenic lines (ANK-1A to -1D); a red-grained line, AUS 1490, and its white-grained mutant line (EMS-AUS). ANK lines showed higher levels of grain dormancy than NS-67 at harvest maturity. AUS 1490 grain also showed higher dormancy than EMS-AUS grain. These results suggest that the R gene for grain colour can enhance grain dormancy. However, the dormancy effect conferred by the R gene was not large, suggesting that it plays a minor role in the development of grain dormancy. Water extracts of AUS 1490 and EMS-AUS bran contained germination inhibitors equivalent to 1-10 microM ABA, although there was no difference in the amount of inhibitors between AUS 1490 and EMS-AUS. Thus, the grain colour gene of AUS 1490 did not appear to enhance the level of grain dormancy by accumulating germination inhibitors in its bran. Sensitivity to ABA of embryos was higher in grains collected around harvest-maturity for ANK lines and AUS 1490, compared with NS-67 and EMS-AUS. The R gene might enhance grain dormancy by increasing the sensitivity of embryos to ABA.     

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

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

Combinational dormancy in seeds of Sicyos angulatus (Cucurbitaceae,tribe Sicyeae)

Seeds with a water‐impermeable seed coat and a physiologically dormant embryo are classified as having combinational dormancy. Seeds of Sicyos angulatus (burcucumber) have been clearly shown to have a water‐impermeable seed coat (physical dormancy [PY]). The primary aim of the present study was to confirm (or not) that physiological dormancy (PD) is also present in seeds of S. angulatus. The highest germination of scarified fresh (38%) and 3‐month dry‐stored (36%) seeds occurred at 35/20°C. The rate (speed) of germination was faster in scarified dry‐stored seeds than in scarified fresh seeds. Removal of the seed coat, but leaving the membrane surrounding the embryo intact, increased germination of both fresh and dry‐stored seeds to > 85% at 35/20°C. Germination (80–100%) of excised embryos (both seed coat and membrane removed) occurred at 15/6, 25/15 and 35/20°C and reached 95–100% after 4 days of incubation at 25/15 and 35/20°C. Dry storage (after‐ripening) caused an increase in the germination percentage of scarified and of decoated seeds at 25/15°C and in both germination percentage and rate of excised embryos at 15/6°C. Eight weeks of cold stratification resulted in a significant increase in the germination of scarified seeds at 25/15 and 35/20°C and of decoated seeds at 15/6 and 25/15°C. Based on the results of our study and on information reported in the literature, we conclude that seeds of S. angulatus not only have PY, but also non‐deep PD, that is, combinational dormancy (PY + PD).