Dormancy-breaking requirements and level of morphophysiological dormancy (MPD) were determined for seeds of Ilex formosana and I. uraiensis from the subtropical region, and seeds of I. rotunda from both the subtropical and tropical regions of Taiwan. We hypothesized that some Ilex species would have deep simple MPD broken by warm stratification. Germination of seeds and embryo growth was monitored at 30/20, 25/15, 20/10, 15/5°C and at 25°C. Seeds were cold-stratified and then incubated at 25/15°C, and seeds treated with GA3 and GA4 were incubated at 25/15°C. Fresh seeds reached 50% germination after 11–45 weeks of warm stratification. Pre-treatment with GA increased germination percentages of I. formosana and I. rotunda (tropical) but not I. uraiensis and I. rotunda (subtropical), while cold stratification did not promote germination of either species but increased the germination rate of I. rotunda (tropical). Embryo length in seeds of all species increased ≥710% prior to root emergence, and growth occurred during warm stratification. The positive response to GA and relatively short time for beginning of germination and to reach 50% germination indicate non-deep simple MPD in seeds of I. formosana and I. rotunda (tropical). The negative response to GA and long time for beginning of germination and to reach 50% germination indicate deep simple MPD in seeds of I. uraiensis and I. rotunda (subtropical). Thus, in both the subtropical and tropical regions of Taiwan, the seeds of Ilex species have non-deep simple and deep simple MPD that are broken by warm stratification. Furthermore, GA treatment increases the germination rate and percentage of Ilex seeds with non-deep simple MPD, and cold stratification promotes the seed germination rate of Ilex species with non-deep simple MPD in tropical region.

Plant community-level studies on seed dormancy traits are important to understand and determine the significance of seed dormancy in different ecosystems. Hence, we studied seed dormancy and other related seed biological traits of 42 selected tropical lowland rainforest tree species from Sri Lanka, aiming to address seed dormancy class(es) for a biodiverse tropical lowland wet zone forest community and the relationship between dormancy classes, forest strata and seed dispersal mechanisms. Seed germination, imbibition, embryo length:seed length ratio, embryo morphology and the effect of gibberrelic acid on seed germination were determined. Sixty-two percent of the species with T50 < 30 days were identified as having fast-germinating seeds and the remaining 38% with T50 > 30 possessed slow-germinating seeds. Seeds of 33 species had fully developed embryos, while nine species had underdeveloped embryos; three had morphological dormancy (MD) and six morphophysiological dormancy (MPD). Treatment with gibberellic acid revealed physiological dormancy (PD) in seeds of six species, and the response to manual scarification confirmed physical dormancy (PY) in seeds of Pericopsis moonina. The majority of tropical lowland rainforests had non-dormant (ND) species (62%), and 14.3, 14.3, 7 and 2.3% of the species had MPD, PD, MD, and PY, respectively. Non-dormancy decreased for taxa from the upper strata to the lower strata of the forest. ND seeds were dispersed during the rainy season. Thus, non-dormancy seems to be the most dominant germination behaviour among the tree species in the lowland rainforest of Sri Lanka with the class of dormancy related to forest strata and dispersal time.

This review provides a revised and expanded word-formula system of whole-seed primary dormancy classification that integrates the scheme of Nikolaeva with that of Baskin and Baskin. Notable changes include the following. (1) The number of named tiers (layers) in the classification hierarchy is increased from three to seven. (2) Formulae are provided for the known kinds of dormancy. (3) Seven subclasses of class morphological dormancy are designated: ‘dust seeds’ of mycoheterotrophs, holoparasites and autotrophs; diaspores of palms; and seeds with cryptogeal germination are new to the system. (4) Level non-deep physiological dormancy (PD) has been divided into two sublevels, each containing three types, and Type 6 is new to the system. (5) Subclass epicotyl PD with two levels, each with three types, has been added to class PD. (6) Level deep (regular) PD is divided into two types. (7) The simple and complex levels of class morphophysiological dormancy (MPD) have been expanded to 12 subclasses, 24 levels and 16 types. (8) Level non-deep simple epicotyl MPD with four types is added to the system. (9) Level deep simple regular epicotyl MPD is divided into four types. (10) Level deep simple double MPD is divided into two types. (11) Seeds with a water-impermeable seed coat in which the embryo-haustorium grows after germination (Canna) has been added to the class combinational dormancy. The hierarchical division of primary seed dormancy into many distinct categories highlights its great diversity and complexity at the whole-seed level, which can be expressed most accurately by dormancy formulae.

As a contribution to understanding the world biogeography of seed dormancy in the cosmopolitan genus Ilex, we studied seeds of I. argentina, I. brasiliensis, I. brevicuspis, I. dumosa, I. paraguariensis and I. theezans from the subtropical region of Argentina. We hypothesized that seeds of these species have non-deep simple morphophysiological dormancy (MPD). Effects of temperature, cold stratification and gibberellic acid (GA3) on seed germination and embryo growth were tested. Regardless of incubation temperature, little or no germination occurred for any species until ≥6 weeks. There was an up to 3-fold increase in embryo length to seed length (E:S) ratio before seeds germinated, and embryos grew only during warm-stratifying conditions. Seeds of I. brasiliensis, I. brevicuspis and I. theezans had non-deep simple MPD and germinated to ≥80% after 12, 24 and 16 weeks, respectively. Cold stratification increased germination of I. brasiliensis and I. brevicuspis, and GA3 increased the rate but not final germination percentage of I. brasiliensis and I. theezans. Fresh seeds of I. dumosa required 40 weeks of warm stratification to germinate to 53%, while those after-ripened for 2 months germinated to 81% after 30 weeks; this species has intermediate simple MPD. Seeds of I. argentina and I. paraguariensis germinated to 15 and 21%, respectively, after 40 weeks of warm stratification and did not after-ripen or respond to GA3; these seeds have deep simple MPD. This is the first report of intermediate and deep simple MPD that is broken by warm stratification, thereby increasing our knowledge of seed dormancy in Ilex and in subtropical regions.

Ferula ovina is a perennial Apiaceae with great medicinal and economic value. This study was conducted to better understand the dormancy and germination behaviour of its seeds. Experiments included temperature requirements for both embryo growth and germination under field and laboratory conditions, the effects of warm stratification, gibberellic acid (GA3), dry after-ripening, exogenous hydrogen peroxide (H2O2) and diphenyleneiodonium (DPI) on dormancy, localizing the accumulation of superoxide (O2−) and measuring endogenous contents of H2O2 in embryos during cold and warm stratification. Embryos were under-developed and did not germinate within one month at temperatures higher than 10°C. Among all treatments, cold stratification and exogenous H2O2 could break dormancy. However, the application of DPI reduced growth and the germination of the embryo at 3°C. During cold stratification, the embryonic axes elongated more than the cotyledons, which coincided with earlier production of O2− in the axes. Only these embryos could eventually complete growth and germinate. Moreover, such asymmetric growth between the axes and cotyledons was also observed when seeds were treated with exogenous H2O2. On the other hand, both axes and cotyledons displayed the same growth over treatments with warm stratification and DPI. Overall, seeds of F. ovina exhibit the characteristics of deep complex morphophysiological dormancy. Moreover, the embryos solely require cold to complete growth and to germinate, and this process seems to be mediated by reactive oxygen species. Under natural conditions, seeds germinate during winter in cold soil, and shoots emerge in spring.

Little information is available on seed dormancy of members of the Zingiberales and especially the Zingiberaceae. Our aim was to investigate the dormancy breaking and germination requirements of Alpinia galanga in vitro with a minimum number of seeds, using the move-along experiment. The mass of imbibed seeds increased by 17.5% in 1 d, showing that seeds were water permeable. The best germination in the move-along experiment (86.7%) was obtained when seeds were exposed to the sequence of temperature regimes that began with winter (20/10°C), and seeds began to germinate after 6 weeks at this temperature regime. Seeds dry stored for 4 months and then incubated at the sequence of temperature regimes that began with summer (30/20°C) started germinating in the sixth week at this temperature regime and had germinated to 93.3% after 18 weeks. Seeds kept dry for 4 months and then treated with 50 mg l−1 gibberellic acid (GA3) began to germinate at 30/20°C after 2 weeks. Control seeds incubated continuously at 20/10, 25/15 or 30/20°C germinated to 80.6, 77.8 and 60.0%, respectively. When incubated at 15, 20, 25 or 30°C, the ideal temperature for embryo growth was 20°C. Since GA3 and dry storage can break non-deep physiological dormancy and embryos grew during warm stratification, seeds of A. galanga have non-deep simple morphophysiological dormancy (MPD). This is the first report of non-deep simple MPD in the Zingiberaceae.

The Araliaceae is known to have seeds with underdeveloped embryos that must grow prior to radicle emergence, and thus they have morphological (MD) or morphophysiological (MPD) dormancy. Araliaceae is one of about 15 families with woody species in the tropical montane zone, and in Hawaii 15 species occur in the montane. Our purpose was to determine if seeds of the Hawaiian Araliaceae species Cheirodendron trigynum subsp. trigynum have MD or MPD and, if MPD, what level. In a move-along experiment, some seeds were incubated continuously at 15/6, 20/10 or 25/15°C, while others were moved sequentially from low to high or from high to low temperature regimes. Germination percentages and embryo growth were monitored. Also, the effects of cold and warm stratification on dormancy break were determined. Seeds had physiological dormancy (PD) in addition to small embryos that grew prior to germination, and thus MPD. PD was broken slowly ( ≥ 12 weeks), after which embryos grew rapidly, followed by root and shoot emergence. Embryos grew at temperatures suitable for warm stratification; thus, seeds have Type 1 non-deep simple MPD; the dormancy formula is C1bBb. Seeds from Oahu germinated to 94–100% at 15/6, 20/10 and 25/15°C, while those from the Big Island germinated to high percentages only at 15/6 and 20/10°C. Temperature shifts improved germination of seeds from the Big Island, and movement from either low to high or from high to low temperature regimes was effective in promoting germination. This is the first report of non-deep simple MPD in the Araliaceae.

The three tropical species Gomphandra luzoniensis, Nothapodytes nimmoniana and Goniothalamus amuyon contain important cancer-fighting drugs; however, little is known about how to propagate these species from seeds. Thus, the aim of this study was to determine the germination requirements of seeds of each of these three species in order to provide an effective protocol to produce plants. Fresh seeds of G. luzoniensis germinated up to 73% at high temperatures in light in 4 weeks, and embryos were underdeveloped. Most seeds had morphological dormancy (MD), but a proportion of them had morphophysiological dormancy (MPD). Fresh seeds of N. nimmoniana germinated up to 50% in light in 4 weeks, embryo length increased by 17% before radicle emergence and ≥ 89% of the seeds had germinated after incubation for 6 weeks in light at high temperatures. Thus, about 50% of the seeds have MD and about 50% MPD. Fresh seeds of G. amuyon incubated at 30°C in light for 4 weeks germinated to 69%, whereas at the other incubation temperatures germination took longer than 4 weeks. Embryo length increased 213% before radicle emergence, and after 8 weeks of incubation at high temperatures ≥ 80% of the seeds had germinated. As in the other two species, the seed population consisted of a mixture of MD and MPD. Incubation of seeds of these three species at high temperatures (e.g. 25, 30 and 30/20°C) for up to 2 months is recommended for germination and thus seedling production.

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.

Palm diaspores are reported to have various kinds of dormancy. However, (1) the embryo is underdeveloped; (2) the endocarp is water permeable; and (3) the diaspores take a long time to germinate. Thus, we conclude that the diaspores of the majority of palm species have morphophysiological dormancy (MPD). The ones that do not have MPD are morphologically dormant.

Although it has been speculated that seeds of the gymnosperm family Podocarpaceae have an underdeveloped embryo, no detailed studies have been done to definitively answer this question. Our purpose was to determine if embryos in seeds of two species of Podocarpaceae, Podocarpus costalis and Nageia nagi, from Taiwan are underdeveloped and to examine the kind of dormancy the seeds have. Embryos in fresh seeds of P. costalis were 4.6 ± 0.5 mm long, and they increased in length by about 54% before radicle emergence (germination), demonstrating that the embryo is underdeveloped at seed maturity. Seeds germinated to >90% at 30/20, 25/15 and 25°C in light in ≤ 4 weeks, without any cold stratification pretreatment. Thus, seeds of P. costalis have morphological dormancy (MD). Embryos in fresh seeds of N. nagi were 7.4 ± 0.8 mm long and they increased in length by about 39% before radicle emergence (germination) occurred, indicating that the embryo is underdeveloped at seed maturity. Seeds germinated to < 25% at 30/20 and 25°C in light in 4 weeks but to >90% at the same temperatures in 12 weeks. Thus, most seeds of N. nagi have morphophysiological dormancy (MPD). Although underdeveloped embryos are considered to be a primitive condition in seed plants, they also occur in the most advanced orders. The occurrence of underdeveloped embryos in Podocarpaceae documents that they are not restricted to a basal clade in gymnosperms.

Heretofore, no detailed account was available on seed dormancy and germination of a member of the Colchicaceae (Liliales). Thus, the primary aim of this study was to do an in-depth investigation of the temperature requirements for dormancy break and germination in seeds of Merendera montana (Colchicaceae) at the embryo and whole-seed levels under near-natural temperatures in a non-heated frame shade-house and under controlled conditions in the laboratory. Mean embryo length in fresh seeds was c. 0.57 mm and embryos had to grow to at least 1.30 mm before radicle emergence. Embryos grew to full size and seeds completed germination (radicles emerged) when they were stratified at 28/14°C for 60 d followed by a cool temperature for 60 d and then incubated at a cool temperature for 30 d. The optimum cool stratification temperature for dormancy-break was 10°C. Thus, after the moist pretreatment at 28/14°C+10°C, radicle emergence was>93% at all incubation temperatures (5, 15/4 and 20/7°C). In its natural habitat, M. montana seeds are dispersed in June, the embryo elongates to full size in autumn and radicles emerge from early November to early February. Although the shoot does not emerge until March and April, it is not physiologically dormant. The shoot emerged from 80% of the radicle-emerged seeds in 13 d at 20/7°C without a previous cold pretreatment. Seeds of M. montana have non-deep complex morphophysiological dormancy, C1b1aB-C1a. This is the first study on seeds with complex MPD to show a delay in shoot emergence following root emergence despite the shoot being physiologically non-dormant.

Size-class structure was used to monitor the growth of underdeveloped embryos in two seed cohorts of Aristolochia macrophylla, A. tomentosa and A. californica and the results are related briefly to differences in seed dormancy among and within the three species. The size-class method of presenting embryo growth data allows more insight to be gained into the embryo-level cause(s) of seed dormancy than does the cumulative growth curve. This appears to be the first report on use of size-class structure to monitor growth of underdeveloped embryos.

Seeds of Aconitum napellus subsp. castellanum were physiologically dormant at maturity in early autumn, with underdeveloped embryos. Thus they have morphophysiological dormancy (MPD). Embryos in fresh seeds were on average 1.01 mm long, and they had to grow to 3.60 mm before radicle emergence. Cold stratification at 5°C for 5 months with light enhanced the mean embryo length to 2.73 mm (SE = 0.13) and seed germination to 20%. However, with higher temperatures (15/4, 20/7, 25/10, 28/14 and 32/18°C) embryo growth was small, with no seeds germinating. Optimal germination was achieved after 4 months of cold stratification at 5°C followed by incubation at 20/7°C for 1 month with light, when germination ranged between 70 and 79%, depending on seed age, locality and year of collection. Cold stratification could be substituted by the application of GA3 solution, since mean embryo length in seeds incubated at 25/10°C for 1 month with light was 3.52 mm and the germination was 80%. Since cold stratification was the only requirement for the loss of MPD, the longest embryo growth occurred during this treatment, and GA3 promoted MPD loss, we concluded that A. napellus seeds have intermediate complex MPD. Germination was higher in 4-month stored than in freshly matured seeds. A pronounced variability in germinative patterns at inter-annual and inter-population level was recorded.

Previous studies indicated that seeds of Viburnum odoratissimum had only physiological dormancy (PD), but no measurements of embryos were made during the dormancy-break treatments. Thus, we investigated embryo growth and radicle and cotyledon emergence over a range of temperatures. Seeds have underdeveloped embryos, and their length increased about 300% before radicle emergence. Embryos also had PD, as evidenced by delays in beginning of embryo growth (2–3 weeks) and of germination after embryos were elongated (4 weeks). After radicle emergence, epicotyl emergence was delayed 1–8 weeks, depending on incubation temperature, but cold stratification was not required to break PD of the epicotyl. Unlike seeds of many previously studied Viburnum spp., epicotyls of V. odoratissimum have non-deep, rather than deep, PD. Hence, a new level of MPD called non-deep, simple, epicotyl MPD has been identified.

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.

The germination dynamics of Ilex aquifolium L. were studied in central Spain using a 3-year field sowing experiment. Ten triads of bags (n = 30 seeds per bag) were buried randomly in four different microhabitats: inside the holly woodland, along the forest edge, in open grassland and under isolated shrubs. Bags were harvested annually, and seeds were analysed for seed viability, embryo development changes and germination. Holly seeds did not germinate until the second year. Embryo development and seed germination were faster at the forest edge, with a total germination proportion of 63% of sown seeds. Germination in the holly woodland (48%), under the shrubs (46%) and in grassland (46%) were very similar. Thus, recruitment of Ilex populations in this region is not limited by requirements for germination. After the third year, the seed bank was reduced by 80–90% of the initial seeds, so it can be considered as a short-persistent seed bank. Holly seed behaviour can enhance recruitment in semi-open areas, such as forest edges. Linked to the short persistence of its seed bank, this species can be considered an intermediate strategist between mature forest specialists and early successional species. This work contributes to a deeper knowledge of the germination ecology of I. aquifolium, in an effort to determine the relative importance of germination/ dormancy processes within holly population dynamics.

Studies were conducted to determine if small embryos (i.e. low embryo length:seed length ratio) in mature dwarf seeds (0.2–2 mm) are underdeveloped. In this case, they would grow (inside the seed) prior to germination, and seeds would have morphological or morphophysiological dormancy. Prior to radicle emergence, embryo length in seeds of Drosera anglica (Droseraceae), Campanula americana, Lobelia appendiculata, L. spicata (Campanulaceae) and Sabatia angularis (Gentianaceae) increased 0, 103, 182, 83 and 57%, respectively. Since embryo growth did not occur in seeds of D. anglica prior to germination, embryos, although small, are fully developed; seeds have only physiological dormancy. The underdeveloped embryo in seeds of C. americana has little or no physiological dormancy; thus, seeds have morphological dormancy. On the other hand, underdeveloped embryos in seeds of L. appendiculata, L. spicata and S. angularis are physiologically dormant, and seeds have morphophysiological dormancy. Therefore, since small embryos in dwarf seeds may or may not be underdeveloped, assignment of seeds to a dormancy class requires that studies be done to determine if embryos grow inside the seed before germination can occur. Such information is important in understanding the evolutionary relationship of the different kinds of seed dormancy.