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.

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.

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.

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 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.

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.

Rudimentary, broad and small linear embryos occur among members of the most primitive (basal) extant angiosperms, collectively called the ANITA grade (i.e. Amborella, Nymphaeales and Austrobaileyales). Amborella (rudimentary) and Austrobaileyales (rudimentary in Austrobaileyaceae, Illiciaceae and Schisandraceae and small linear in Trimeniaceae) have kinds of embryos that are known to be underdeveloped; consequently, they must grow inside the seed prior to radicle emergence (germination). On the other hand, it is not known if broad embryos need to grow before radicles can emerge, and whether they are underdeveloped or fully developed. Thus, we addressed the question: ‘Is the broad embryo of Nymphaeales also underdeveloped?’. Although the embryo length : seed length ratios in Nymphaea Albert Greenburg, N. capensis var. zanzibariensis and N. immutabilis were 0.311, 0.349 and 0.234, respectively, embryos did not grow prior to radicle emergence. Thus, they are fully developed at seed maturity. If Amborella and Nymphaeales are equally the most basal angiosperms, as some molecular phylogenetic studies indicate, then we must conclude that the broad and rudimentary embryos are equally primitive.

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.

There are two ideas regarding changes in the physiological and ecological tolerances and requirements within plant lineages through geological time. One is that these attributes have changed very little, or not at all (trait stasis), and the other is that they have undergone considerable change (adaptation), as plants shifted to new climatic and vegetation zones. We tested these ideas for seed dormancy-breaking and germination requirements of four species in a subclade of Aristolochia subgenus Siphisia: the three temperate species, A. macrophylla and A. tomentosa (the basal species in the subclade) of eastern USA and A. manshuriensis of East Asia, and the Mediterranean-climate species A. californica endemic to California, USA. A long period at cold-stratifying temperatures was required for growth of the underdeveloped embryo and seed germination in A. californica, whereas embryos grew and seeds germinated in the other three species at warm temperatures, either before or after they were cold stratified. Thus, seeds of A. californica have either intermediate or deep complex morphophysiological dormancy (MPD), whereas those of the three temperate species have either morphological dormancy or non-deep simple MPD. Further, there were quantitative differences in temperature requirements for dormancy-break and germination between the Appalachian A. macrophylla, which did not differ from its sister species A. manshuriensis, and the lowland A. tomentosa. Thus, within this lineage there has been both trait stasis and divergence (adaptation) in the physiology and ecology of seed dormancy and germination.

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.

The effects of temperature on embryo growth, radicle emergence and cotyledon emergence of Gagea lutea (Liliaceae), a perennial herb widely distributed in Europe, eastern Siberia, the Kurile Islands, Sakhalin and the Far East, were monitored outdoors and in laboratory tests. In Japan, this species inhabits open secondary grasslands and deciduous forests. Seeds with an underdeveloped embryo are dispersed in late May/early June in Hokkaido. The embryo elongates in autumn, and the radicle emerges from the seed in mid-October to mid-November, at temperatures of about 15/4°C. However, cotyledons do not emerge until April, after seeds with an emerged radicle are covered with snow (near 0°C) for about 4 months. In laboratory experiments, temperatures of 25/15°C or 20/10°C followed by 5–10°C were required for embryo growth and radicle emergence. Rate and percentage of cotyledon emergence were promoted by keeping seeds with an emerged radicle under snow. The optimum temperature for cotyledon emergence after 81 d under snow was 15/5°C. Thus, G. lutea has deep simple epicotyl morphophysiological dormancy, and this is the first report of epicotyl dormancy in the genus.