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.

Viburnum is a temperate-zone genus that also occurs in mountains of South America and Malesia, and seeds of many species have morphophysiological dormancy (MPD). Information on the level of MPD in seeds of species in various clades of Viburnum potentially would increase our understanding of the evolutionary relationships between the nine levels of MPD. Our aim was to determine the level of MPD in seeds of Viburnum plicatum var. formosanum that is endemic to mountains (1800–3000 m a.s.l.) in Taiwan and a member of the Lutescentia clade. The temperature requirements for embryo growth and root and shoot emergence and response of seeds to gibberellic acid (GA) were determined. No fresh seeds germinated during 16 weeks of incubation at 15/5, 20/10, 25/15, 30/20 or 25°C. Embryo growth and root emergence occurred during moist cold stratification at 5°C or at a temperature sequence of 15/5 to 5°C. During cold stratification, embryos length increased from 0.76 ± 0.06 to 3.40 ± 0.26 mm and the embryo length:seed length ratio from 0.20 ± 0.02 to 0.68 ± 0.07. In a temperature sequence simulating field conditions, embryos grew inside seeds at 5°C, roots emerged at 15/5°C and shoots emerged at 20/10°C. The optimum temperature for embryo growth was 5°C. Neither GA3 nor GA4 was effective in promoting root emergence. We conclude that seeds of V. plicatum var. formosanum have deep complex MPD, which is a first report for Viburnum. Dormancy release during the cool season at high elevations helps to ensure that seeds germinate at the beginning of the warm season.

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.

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.

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.

Although mature seeds of the monogeneric conifer family Cephalotaxaceae sensu stricto have underdeveloped embryos, no definitive studies have been done to classify dormancy in this family. Our primary purpose was to determine the kind of dormancy in seeds of Cephalotaxus wilsoniana and to put the results into a broad phylogenetic context for gymnosperms. The species is of horticultural and medicinal value, and information is needed on how to propagate it efficiently from seeds. Embryo growth and germination were monitored for seeds at warm, cold and warm plus cold temperatures, and germination was monitored for seeds subjected to: (1) cold →  warm →  cold →  warm; and (2) warm →  cold →  warm →  cold →  warm temperature sequences. The effects of gibberellic acids GA3 and GA4 were tested on radicle emergence in ungerminated seeds and on shoot emergence in root-emerged seeds. Germination was promoted by ≥ 36 weeks of warm stratification followed by ≥ 8 weeks of cold stratification, but only if seeds were returned to high temperatures. The underdeveloped embryo must increase in length by >120% before the radicle emerges. Neither GA3 nor GA4 was effective in promoting radicle emergence; however, both plant growth regulators increased rate (but not percentage) of shoot emergence in root-emerged seeds. We conclude that seeds of C. wilsoniana have the deep simple level of morphophysiological dormancy (MPD), C1b-C3-B1b; thus, warm stratification followed by cold stratification and then warm-temperature incubation are required for germination. In gymnosperms, MPD is known in cycads, Ginkgo and now in three families of conifers.

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.

Intact seeds (seed+endocarp) from freshly harvested fruits of Prunus campanulata were dormant, and required 4–6 weeks of warm followed by 8 weeks of cold stratification for maximum germination percentage. Removing both endocarp and seed coat, however, promoted germination in a high percentage of non-stratified seeds. Treatment of intact, non-stratified seeds with gibberellic acid (GA3) was only partially effective in breaking dormancy. However, GA3 promoted germination of non-stratified seeds in which the endocarp (but not the seed coat) had been removed. The order of abscisic acid (ABA) concentration in fresh seeds was endocarp > seed coat > embryo, and its concentration in endocarp plus seed coat was about 6.2-fold higher than that in the embryo. Total ABA contents of seeds subjected to warm and/or cold moist stratification were reduced 6- to 12-fold. A higher concentration of GA4 was detected in embryos of non-dormant than in those of dormant seeds. Fluridone, a carotenoid biosynthesis inhibitor, was efficient in breaking dormancy of Prunus seeds. Paclobutrazol, a GA biosynthesis inhibitor, completely inhibited seed germination, and the inhibitory effect could be partially reversed by GA4, but not by GA3. Thus, dormancy in P. campanulata seeds is imposed by the covering layers. Dormancy break is accompanied by a decrease in ABA content of the covering layers and germination by an increase of embryonic GA4 content.

The embryo length/seed length (E/S) ratio of the early diverging eudicot Trochodendron aralioides is 0.34. Embryos in fresh seeds were 0.36±0.01 mm long, and they increased in length by about 250% (in 20 d) before radicle emergence (germination) occurred, demonstrating that the embryo is underdeveloped at seed maturity. Seeds germinated to 95–100% at 20/10, 25/15 and 30/15°C in light in ≤4 weeks, without any pretreatment, but no seeds germinated in darkness. Thus, seeds of T. aralioides have morphological dormancy (MD), which is considered to be the primitive condition in seed plants, and MD probably has existed in the genus Trochodendron since its origin in the early Tertiary.