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The daffodil Narcissus pseudonarcissus L. contains alkaloids of pharmaceutical interest. Wild daffodil populations have diverse genetic backgrounds and various genetic traits of possible importance. Developing protocols for plant production from seeds may ensure the availability of a large reservoir of individuals as well as being important for species with bulbs that are difficult to acquire. The closely related Narcissus pseudonarcissus subsp. munozii-garmendiae and subsp. nevadensis were investigated in this study because the alkaloids isolated from both are of high pharmacological interest. At the dispersal time, the seeds of both were dormant with underdeveloped embryos, i.e., morphophysiological dormancy (MPD). Experiments were conducted outdoors and under controlled laboratory conditions. Embryo growth and the percentages of radicle and seedling emergence were calculated under different temperature–light stratifications. In N. munozii-garmendiae, embryo growth occurred during warm stratification (28/14 °C or 25/10 °C) and the radicle then emerged when the temperature decreased, but the shoot was dormant. In N. nevadensis, the seeds germinated when cold stratified (5 °C) and then incubated at cool temperatures. Thus, N. munozii-garmendiae and N. nevadensis exhibit different levels of MPD, i.e., deep simple epicotyl and intermediate complex, respectively. Plant production protocols from seeds were established for both taxa in this study.

Background To exploit the ornamental and medicinal purposes of Lonicera harae Makino, L. subsessilis Rehder, L. praeflorens Batalin, and L. insularis Nakai , native to Korea, it is necessary to understand their seed ecology for propagation. In this study, we investigated the seed dormancy type and germination characteristics of seeds of the four Korean native Lonicera species. Results The seeds of the four Lonicera species imbibed water readily, suggesting that the species do not have physical dormancy. Furthermore, the seeds exhibited underdeveloped embryos with only about 15–25% of the length of the seeds at dispersal. The embryos grew to the critical length with approximately 50–80% of the length of the seeds’ development before radicle protrusion. Further, 94.4% and 61.1% of freshly matured seeds of L. insularis and L. harae germinated within 4 weeks after sowing at 15 °C and 20 °C, respectively. Contrarily, L. praeflorens and L. subsessilis seeds did not germinate within 4 weeks under all temperature treatments. At 15 °C, L. praeflorens seeds started to germinate from 5 weeks and the final germination rate was 51.1% at 13 weeks. At 15 °C, L. subsessilis seeds started to germinate from 5 weeks after sowing and the final germination rate was 85.6% at 17 weeks after sowing. Embryo growth and germination of L. praeflorens and L. subsessilis occurred at a relatively high temperature (≥ 15 °C). Conclusions Overall, L. insularis seeds have only morphological dormancy. The seeds of L. harae have approximately 60% and 40% of morphological dormancy and morphophysiological dormancy, respectively. Contrarily, L. praeflorens and L. subsessilis exhibited non-deep simple-type morphophysiological dormancy that requires relatively high temperature (≥ 15 °C) for embryo growth and dormancy breaking. The optimum temperature for the germination of seeds of L. insularis , L. harae , L. praeflorens , and L. subsessilis was 15 °C, 20 °C, 15 °C, and 20 °C, respectively. There was interspecific variation in seed dormancy and germination patterns in the four Lonicera species. The difference in these characteristics within the four Lonicera species could be useful for understanding the seed ecophysiological mechanisms of Lonicera species.

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.

The genus Paeonia (Paeoniaceae) includes many popular ornamentals, has colorful flowers and contains several Chinese medicinal species. The germination protocol for seeds of Paeonia species is complex and impedes the breeding of new cultivars and contributes to the rarity and high cost of the plants. Although numerous reports on seed dormancy/germination in peonies are scattered throughout the literature, most of them are in Chinese. The primary aims of this paper are to provide a general overview of the available information on seed dormancy/germination in peonies and to make some suggestions regarding propagation for the peony industry and breeders. Most Paeonia species have epicotyl dormancy. The embryo is differentiated into organs, but it is underdeveloped (small) and must grow inside the seed before the radicle can emerge. Germination of peony seeds requires warm stratification for embryo growth and radicle protrusion followed by cold stratification for epicotyl growth. In addition, the epicotyl is sensitive to cold stratification only after the root has grown to a certain length. GA₃ treatment enhances embryo growth and subsequent germination percentages. Further investigations on the physiology, genetics and proteomics would contribute to a better understanding of seed dormancy in Paeonia.

Background Panax notoginseng (Burk) F.H. Chen is an essential plant in the family of Araliaceae. Its seeds are classified as a type of morphophysiological dormancy (MPD), and are characterized by recalcitrance during the after-ripening process. However, it is not clear about the molecular mechanism on the after-ripening in recalcitrant seeds. Results In this study, exogenous supply of gibberellic acid (GA 3 ) with different concentrations shortened after-ripening process and promoted the germination of P. notoginseng seeds. Among the identified plant hormone metabolites, exogenous GA 3 results in an increased level of endogenous hormone GA 3 through permeation. A total of 2971 and 9827 differentially expressed genes (DEGs) were identified in response to 50 mg L −1 GA 3 (LG) and 500 mg L −1 GA 3 (HG) treatment, respectively, and the plant hormone signal and related metabolic pathways regulated by GA 3 was significantly enriched. Weighted gene co-expression network analysis (WGCNA) revealed that GA 3 treatment enhances GA biosynthesis and accumulation, while inhibiting the gene expression related to ABA signal transduction. This effect was associated with higher expression of crucial seed embryo development and cell wall loosening genes, Leafy Contyledon1 ( LEC1 ), Late Embryogenesis Abundant ( LEA ), expansins ( EXP ) and Pectinesterase ( PME ). Conclusions Exogenous GA 3 application promotes germination and shorts the after-ripening process of P. notoginseng seeds by increasing GA 3 contents through permeation. Furthermore, the altered ratio of GA and ABA contributes to the development of the embryo, breaks the mechanical constraints of the seed coat and promotes the protrusion of the radicle in recalcitrant P. notoginseng seeds. These findings improve our knowledge of the contribution of GA to regulating the dormancy of MPD seeds during the after-ripening process, and provide new theoretical guidance for the application of recalcitrant seeds in agricultural production and storage.

Available information on seed dormancy for various members of the Saxifragales and phylogenetic relationships within this order allowed us to accurately predict that Daphniphyllum glaucescens seeds have morphophysiological dormancy (MPD). However, our hypothesis that seeds had deep simple epicotyl MPD, i.e., nondeep physiological dormancy (PD) in root and deep PD in shoot, was not supported. Both the root and the shoot (cotyledons) of the underdeveloped embryo of D. glaucescens have nondeep PD. Exposure to moderate (15°/6°, 20°/10°C), rather than high (25°/15°C), temperatures for 10-12 wk broke the PD of the hypocotyl/root. After hypocotyl emergence, seeds with an attached developing root system did not require cold stratification to break the PD of the shoot. The PD of the shoot was broken by an additional 10-12 wk at moderate temperatures, during which time cotyledons slowly grew inside the seed. As cotyledons grew, all of the hypocotyl was pushed out of the seed, and the final cotyledon length was almost twice that of the seed; at this point, the folded cotyledons emerged. Since the level of PD in root and shoot may or may not be the same, we advocate stating the level of PD in both root and shoot when epicotyl MPD is described in a species. Thus, seeds of D. glaucescens have nondeep simple (root)-nondeep simple (epicotyl) MPD, which is written as C1bB(root)-C1bB(shoot) in the formula system of Nikolaeva. This is the first report of this level of epicotyl MPD in the Saxifragales.

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.

Background Seed dormancy and germination are key components of plant regeneration strategies. Aconitum barbatum is a plant commonly found in northeast China. Although it has potential for use in gardening and landscaping, its seed dormancy and regeneration strategy, which adapt to its natural habitat, are not well understood. Our aim was to identify conditions for breaking A. barbatum seed dormancy and determine its dormancy type. Embryo growth and germination were determined by collecting seeds over time in the field. Laboratory experiments that control light, temperature, and stratification period were conducted to assess dormancy breaking and germination, and GA 3 was used to identify dormancy type. Results Seeds of A. barbatum have undeveloped embryos with physiological dormancy at maturity in autumn. The embryo-to-seed length ratio increases from 0.33 to 0.78 before the emergence of the radical. Under natural environmental conditions, embryo development begins in early winter. Laboratory experiments have shown that long-term incubation under 4 °C (cold stratification) promotes embryo development and seed dormancy break. With an extension of cold stratification, an increase in germination percentages was observed when seeds were transferred from 4 °C to warmer temperatures. Seeds exposed to light during incubation show a higher germination percentage than those kept in the dark. Seed germination can also be enhanced by a 100 mg/L GA 3 concentration. Conclusions Seeds of A. barbatum display intermediate complex morphophysiological dormancy at maturity. In addition to the underdeveloped embryo, there are also physiological barriers that prevent the embryo from germinating. Dormancy breaking of A. barbatum seeds can be achieved by natural winter cold stratification, allowing seeds to germinate and sprout seedlings at the beginning of the following growing season. Our findings provide valuable insights into the seed dormancy and regeneration strategy of A. barbatum , which could facilitate its effective utilization in gardening and landscaping.

Abstract Background and Aims Substantial evidence supports the hypothesis that morphophysiological dormancy (MPD) is the basal kind of seed dormancy in the angiosperms. However, only physiological dormancy (PD) is reported in seeds of the ANA-grade genus Nymphaea. The primary aim of this study was to determine the kind of dormancy in seeds of six species of Nymphaea from the wet–dry tropics of Australia. Methods The effects of temperature, light and germination stimulants on germination were tested on multiple collections of seeds of N. immutabilis, N. lukei, N. macrosperma, N. ondinea, N. pubescens and N. violacea. Embryo growth prior to hypocotyl emergence was monitored. Key Results Germination was generally <10 % after 28 d in control treatments. Germination percentage was highest at 30 or 35 °C for seeds exposed to light and treated with ethylene or in anoxic conditions in sealed vials of water, and it differed significantly between collections of N. lukei, N. macrosperma and N. violacea. Seeds of N. pubescens did not germinate under any of the conditions. Embryo growth (8–37 % in length) occurred before hypocotyl emergence (germination) in seeds of the five species that germinated. Conclusions Fresh seeds were dormant, and the amount of pregermination embryo growth in seeds of N. lukei and N. immutabilis was relatively small, while in seeds of N. macrosperma, N. ondinea and N. violacea it was relatively large. Thus, seeds of N. lukei and N. immutabilis had PD and those of N. macrosperma, N. ondinea and N. violacea had MPD. Overall, we found that seeds in the most phylogenetically derived clades within Nymphaea have MPD, suggesting that PD is the most likely basal trait within the Nymphaeales. This study also highlights the broad range of dormancy types and germination strategies in the ANA-grade angiosperms.

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 GA 3 and GA 4 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.