Morphologic and cytotaxonomic evaluation of Lomatium tuberosum (Apiaceae)

Lomatium tuberosum Hoover, a south-central Washington endemic, has been considered to be closely related to three other normally purple-flowered lomatiums:L. columbianum, L. cuspidatum, andL. minus. Recent morphologic studies confirm the close phenetic relationship of these four species. Chromosome counts from a purple-flowered population in the eastern range ofL. tuberosum indicaten=11, the same chromosome number earlier determined with yellow-flowered plants from the type locality.     

A new species of Lomatium (Apiaceae) from Utah

A new species ofLomatium belonging to theCynomarathrum group is described and its relationships to other species in the group are indicated.Lomatium junceum appears to be restricted to Emery Co., Utah, on barren clay and shale slopes in the San Rafael Swell and adjacent eastern base of the Wasatch Plateau.     

Deep and intermediate complex morphophysiological dormancy in seeds of Ferula gummosa (Apiaceae)

We tested the hypothesis that seeds of the monocarpic perennial Ferula gummosa from the Mediterranean area and central Asia have deep complex morphophysiological dormancy. We determined the water permeability of seeds, embryo morphology, temperature requirements for embryo growth and seed germination and responses of seeds to warm and cold stratification and to different concentrations of GA₃. The embryo has differentiated organs, but it is small (underdeveloped) and must grow inside the seed, reaching a critical embryo length, seed length ratio of 0.65–0.7, before the seed can germinate. Seeds required 9 weeks of cold stratification at <10°C for embryo growth, dormancy break and germination to occur. Thus, seeds have morphophysiological dormancy (MPD). Furthermore, GA₃ improved the germination percentage and rate at 5°C and promoted 20 and 5% germination of seeds incubated at 15 and 20°C, respectively. Thus, about 20% of the seeds had intermediate complex MPD. For the other seeds in the seed lot, cold stratification (5°C) was the only requirement for dormancy break and germination and GA₃ could not substitute for cold stratification. Thus, about 80% of the seeds had deep complex MPD.     

Dispersal limitation inferred from an experimental translocation of Lomatium (Apiaceae) species outside their geographic ranges

Determining limitations on poleward range expansion is important for predicting how climate change will alter the distribution of species. For most species, it is not known what factors set their distributional limits and the role dispersal limitation might play if range‐limiting factors were altered. We conducted a transplant study of three related and co‐occurring Lomatium species at their northern range limits to test competing hypotheses of range limitation. We added seeds to experimental plots inside and outside the species’ geographic range (a regional treatment) in a replicated design with vegetation intact and vegetation reduced (a disturbance treatment) and with herbivore access and herbivore exclusion (an herbivory treatment). Germination and reemergence were measured through two growing seasons, along with community‐level variables. A fully‐crossed linear mixed model revealed that Lomatium survivorship outside the current range was as good or better than survivorship within the range, at least when the vegetative community remained intact. This suggests that the species are dispersal limited. Germination often was improved in the presence of an intact vegetative community, but this potentially facilitative effect was absent in second‐year reemergence. Plots exposed to herbivory had slightly, but significantly, reduced germination, though reemergence did not differ between herbivore treatments. Lomatium dissectum, a rare species, had significantly lower survivorship than its congeners, suggesting that range shifts in rare taxa may be particularly difficult. Seed additions beyond species’ range limits may be a strategy for overcoming dispersal limitation and assisting species in poleward migrations.     

Molecular mechanisms and hormonal regulation underpinning morphological dormancy: a case study using Apium graveolens (Apiaceae)

Underdeveloped (small) embryos embedded in abundant endosperm tissue, and thus having morphological dormancy (MD) or morphophysiological dormancy (MPD), are considered to be the ancestral state in seed dormancy evolution. This trait is retained in the Apiaceae family, which provides excellent model systems for investigating the underpinning mechanisms. We investigated Apium graveolens (celery) MD by combined innovative imaging and embryo growth assays with the quantification of hormone metabolism, as well as the analysis of hormone and cell-wall related gene expression. The integrated experimental results demonstrated that embryo growth occurred inside imbibed celery fruits in association with endosperm degradation, and that a critical embryo size was required for radicle emergence. The regulation of these processes depends on gene expression leading to gibberellin and indole-3-acetic acid (IAA) production by the embryo and on crosstalk between the fruit compartments. ABA degradation associated with distinct spatiotemporal patterns in ABA sensitivity control embryo growth, endosperm breakdown and radicle emergence. This complex interaction between gibberellins, IAA and ABA metabolism, and changes in the tissue-specific sensitivities to these hormones is distinct from non-MD seeds. We conclude that the embryo growth to reach the critical size and the associated endosperm breakdown inside MD fruits constitute a unique germination programme.     

Epicotyl morphophysiological dormancy in seeds of Lilium polyphyllum (Liliaceae)

Dormancy-breaking and seed germination studies in genus Lilium reveal that the majority of Lilium spp. studied have an underdeveloped embryo at maturity, which grows inside the seed before the radicle emerges. Additionally, the embryo, radicle or cotyledon has a physiological component of dormancy; thus, Lilium seeds have morphophysiological dormancy (MPD). A previous study suggested that seeds of Lilium polyphyllum have MPD but the study did not investigate the development of the embryo, which is one of the main criteria to determine MPD in seeds. To test this hypothesis, we investigated embryo growth and emergence of radicles and epicotyls in seeds over a range of temperatures. At maturity, seeds had underdeveloped embryos which developed fully at warm temperature within 6 weeks. Immediately after embryo growth, radicles also emerged at warm temperatures. However, epicotyls failed to emerge soon after radicle emergence. Epicotyls emerged from >90% seeds with an emerged radicle only after they were subjected to 2 weeks of cold moist stratification. The overall temperature requirements for dormancy-breaking and seed germination indicate a non-deep simple epicotyl MPD in L. polyphyllum.     

A possible mechanism of high temperature dormancy regulation in seeds of Avena sativa L. (cv. Moyencourt)

Embryos of Avena sativa L. (cv. Moyencourt) show no high temperature dormancy. The dormancy is induced by the presence of endosperm-aleurone part of the seed. Germination of isolated embryos at 30°C can be prevented by ABA and the inhibition is reversed by GA. Inhibitors of GA synthesis also inhibit embryo germination. The embryos of dormant and non-dormant seeds vary greatly in their sensitivity to exogenous ABA. High temperature dormancy of the entire seeds can be relieved by low concentrations of ethanol. On the basis of these facts a hypothetic model is proposed showing how interaction between endogenous GA and ABA-like inhibitory substance, may regulate the high temperature dormancy of the seeds.     

A quantitative model for loss of primary dormancy and induction of secondary dormancy in imbibed seeds of Orobanche spp

Seeds of three species of Orobanche were conditioned (e.e. stored fully imbibed in darkness) for periods up to 210 d in order to model relief of primary dormancy and induction of secondary dormancy. The data were consistent with the hypothesis that periods for loss and induction of formancy and loss of viability are normally distributed in populations of imbibed sees and that these three processes are independent. There was a positive, linear relationship between the rate of loss of primary dormancy and temperature from 10-30C in O. aegyptiaca and O. cernua and 10-25C in O. crenata. In all three species, the rate of induction of secondary dormancy was highest at 10°C and decreased with increase of temperature up to about 20°C, above which there was little further change in the rate with temperature. The resulting model explained over 90% of the variation in germination after conditioning in both O. aegyptiaca and O. cernua. In O. crenata, however, this model was only satisfactory at 10 and 15°C. At higher temperatures, dormancy was relatively stable for periods of conditioning from 70 to about 154 d. Possible explanations for this are discussed. Applications of these models for estimating the time required to reduce Orobanche infestations in the field are also briefly discussed.Key words: Orobanche spp., broomrape, seed dormancy, temperature, dormancy model      

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).     

Morphophysiological dormancy in seeds of the ANA grade angiosperm Schisandra arisanensis (Schisandraceae)

We determined the kind of seed dormancy in Schisandra arisanensis, an ANA grade ([A]mborellales [N]ymphaeales [A]ustrobaileyales) angiosperm with medicinal value. Seeds have small underdeveloped embryos, and following seed maturity their length increased approximately 360% before radicle emergence. Germination was delayed 6–8 weeks, and the percentage and rate were much higher at 15/6, 20/10 and 25/15°C than at 30/20°C. For seeds incubated at 5/5°C (8 weeks) → 15/6°C (4 weeks) → 20/10°C (8 weeks) → 25/15°C (12 weeks) → 20/10°C (5 weeks), embryos grew at 15/6°C → 20/10°C, and almost all seeds that germinated (89%) did so at 20/10°C → 25/15°C. When seeds were incubated in a complementary temperature sequence, 25/15°C (12 weeks) → 20/10°C (8 weeks) → 15/6°C (4 weeks) → 5/5°C (9 weeks) → 15/6°C (4 weeks), embryos grew at 25/15°C → 20/10°C. Nearly all seeds that germinated (93%) did so at 25/15°C → 20/10°C and at 15/6°C following 9 weeks at 5/5°C. Based on the temperature requirements for embryo growth and seed germination, seeds of this species have non‐deep simple morphophysiological dormancy (C_1bB).     

Transition from primary dormancy to secondary dormancy in cocklebur seeds

Abstract The transition from primary dormancy to secondary dormancy was examined using upper cocklebur (Xanthium pennsylvanicum Wallr.) seeds. The non-after-ripened seeds with primary dormancy responded to chilling, anoxia, KCN, and NaN₃ with an increase in germination. However, their maximal responses to these treatments only occurred after a period of water imbibition, probably a reflection of the increasing growth potential of the axial tissue which was accompanied by the increase in the capacities of respiration and ethylene production. On the other hand, the establishment of secondary dormancy was accompanied by a decrease in respiration and ethylene production of seeds, and in the growth potential of both axial and cotyledonary tissues. The decrease in growth potential of these tissues occurred regardless of whether they were excised from after-ripened seeds or non-after-ripened seeds. It is inferred that the primary dormancy of cocklebur seeds is a state maintained in un-germinated seeds for a long time through a spontaneous transition to secondary dormancy.     

Endosperm dormancy breakage in olive seeds

Seeds of Olea europaea L. ssp. oleaster Hoffm. and Link freed from the sclerous endoearp and incubated in water at 15 or 25°C in darkness or in 12:12 h white light:dark conditions, did not germinate, due to dormancy imposed by the endosperm. Seeds also did not germinate when incubated in abscisic acid, gibberellic acid, kinetin or zeatin in darkness and at cither 15 or 25°C. SAN 9789 |4-chloro-5-(methylamine)-2-(a,a,a-trifluoro-m-tolyl)-3-(2H)-pyridazmone] did not promote germination at 15°C but it did to a 75% level at 25°C. This promoting effect of SAN was counteracted by abscisic acid. Cultures of naked embryos grew equally well in the presence or absence of SAN 9789. 6-Benzylaminopurine promoted whole seed germination to a 15% level.     

Seasonal regulation of dormancy in Chrysopa carnea (Neuroptera)

In Ithaca, New York, diapause incidence in Chrysopa carnea exceeds 50 per cent among adults emerging between 31 August and 3 September. Apparently, decreasing late summer day lengths, which exceed the stationary critical photoperiod in the laboratory, act on the sensitive stages and induce diapause. Full diapause intensity occurs about 2 weeks after adult emergence; complete winter coloration appears approximately 2 weeks later. Photoperiodic stimili in combination with the geographic strain largely determine diapause depth.Decreasing autumn day lengths decelerate diapause development and maintain diapause until the winter solstice. Subsequently, short day lengths and cold temperatures slow diapause development. In nature, neither long days, increasing day lengths, nor chilling hasten diapause termination. Diapause duration is largely determined by the interaction between inducing photoperiod, maintaining photoperiodic and thermal stimuli, and geographic strain. In Ithaca, diapause ends (all response to photoperiod ceases) between 22 January and 12 March.After diapause ends, the animals retain their diapause characteristics until temperatures exceed 4°C. Post-diapause reproductive development in females requires approximately 100 heat units above 4°C; however, below 8°C mating and oviposition are absent. Although the winter coloration remains, mating occurs within 1 day after transfer from 4 to 24°C. Temperature determines the rate at which the green summer colour returns.     

Metabolome profiling of stratified seeds provides insight into the regulation of dormancy in Davidia involucrata

Dove tree (Davidia involucrata), a tertiary vestige species, is well-adapted to cool conditions. Dormancy in D. involucrata seed lasts for an extremely long period of time, typically between 3 and 4 years, and this characteristic makes the species an excellent model for studying the mechanisms of seed dormancy. The molecular mechanisms governing germination control in D. involucrata are still unknown. Seed stratification have been reported to enhance germination in recalcitrant seeds. We performed a widely targeted metabolome profiling to identify metabolites and associated pathways in D. involucrata seeds from six different moist sand stratification durations (0–30 months) using the ultra-high-performance liquid chromatography-Q Exactive Orbitrap-Mass spectrometry. There was an increasing germination rate with prolonged stratification durations (12–30 months). Furthermore, we detected 10,008 metabolites in the stratified seeds. We also detected 48 differentially accumulated metabolites (DAMs) between all stratification periods in the seeds, with 10 highly conserved metabolites. Most of the differentially accumulated metabolites between unstratified and stratified seeds were enriched in purine metabolism, pyrimidine metabolism, flavone and flavonol biosynthesis, phenylpropanoid biosynthesis, and arginine biosynthesis pathways. Key phytohormones, abscisic acid, indole-3 acetic acid, and sinapic acid were differentially accumulated in the seeds and are predicted to regulate dormancy in D. involucrata. We have provided extensive metabolic information useful for future works on dove tree germination study.     

A proposed mechanism for physical dormancy break in seeds of Ipomoea lacunosa (Convolvulaceae)

Background and Aims

The water-impermeable seeds of Ipomoea lacunosa undergo sensitivity cycling to dormancy breaking treatment, and slits are formed around bulges adjacent to the micropyle during dormancy break, i.e. the water gap opens. The primary aim of this research was to identify the mechanism of slit formation in seeds of this species.

Methods

Sensitive seeds were incubated at various combinations of relative humidity (RH) and temperature after blocking the hilar area in different places. Increase in seed mass was measured before and after incubation. Scanning electron microscopy (SEM) and staining of insensitive and sensitive seeds were carried out to characterize these states morphologically and anatomically. Water absorption was monitored at 35 and 25 °C at 100 % RH.

Key Results

There was a significant relationship between incubation temperature and RH with percentage seed dormancy break. Sensitive seeds absorbed water vapour, but insensitive seeds did not. Different amounts of water were absorbed by seeds with different blocking treatments. There was a significant relationship between dormancy break and the amount of water absorbed during incubation.

Conclusions

Water vapour seals openings that allow it to escape from seeds and causes pressure to develop below the bulge, thereby causing slits to form. A model for the mechanism of formation of slits (physical dormancy break) is proposed.Key words: Convolvulaceae, Ipomoea lacunosa, dormancy-breaking mechanism, physical dormancy, seeds, sensitivity cycling, water vapour     

Environmental and hormonal regulation of seed dormancy and germination in Cape fynbos Leucospermum R.Br. (Proteaceae) species

The endogenous levels of abscisic acid (ABA), cytokinins (CKs) and gibberellins (GA₁/GA₃ combined) in Leucospermum glabrum embryos were monitored in axes and cotyledons separately during normal germination. Plant growth substance changes were correlated with known morphological, structural and ultrastructural events in the embryo of Proteaceae. The effect of exogenous application of 6-benzyladenine (BA) and GA₄₊₇ under three known dormancy-enforcing environmental conditions were studied in L. glabrum and L. cordifolium. The endogenous levels of the hormone classes GAs and CKs changed phasically during normal germination under a single alternating temperature regime. GA₁/GA₃ levels increased in cotyledons within 3 d of hydration while at the same time initial CK levels decreased. Following this transient peak GAs fell to a low level throughout the germinative period. Subsequently the CKs, Z and ZR, and to a lesser extent their dihydro-derivatives, appeared in both the axes and the cotyledons as fluctuating, transient peaks. Early increases in GAs are thought to control the induction of the germination process. The CK pattern suggests that CKs control at least three major processes of germination sensu stricto following induction: 1) early mobilization of protein and lipid reserves in the axis and later in cotyledons, 2) cotyledon expansion which causes the endotesta to split permitting radicle protrusion and 3) later, radicle growth.Our results indicate that dormancy in intact Leucospermum seeds is enforced by embryo anoxia, regulated by the impermeable exotesta. In addition synthesis of or tissue sensitizing to both hormone classes GAs and CKs depends on moderately low temperature as the primary environmental requirement. For GA synthesis a secondary, daily pulse of high temperature is required. Inhibitory hormones, specifically ABA, appear not to play a role.Abbreviations ABA Abscisic acid - BA 6-benzyladenine - CK Cytokinin - DHZ Dihydrozeatin - DHZR Dihydrozeatin riboside - GA Gibberellin - HPLC High performance liquid chromatography - iP Isopentenyladenine - IPA Isopentenyladenosine - PGS Plant growth substance - RIA Radioimmunoassay - Z Zeatin - ZR Zeatin riboside     

Morphological dormancy in seeds of the autumn-germinating shrub Lonicera caerulea var. emphyllocalyx (Caprifoliaceae)

To better understand the germination ecophysiology of the genus Lonicera , the dormancy class, temperature requirements for embryo growth and radicle emergence and phenology of seedling emergence were determined for Lonicera caerulea var. emphyllocalyx . At maturity, seeds have an underdeveloped embryo (approximately 28% of the length of full-grown embryos). Embryos in fresh seeds grew to full length at 15, 20, 20/10 and 25/15°C within 3 weeks, but failed to grow at ≤ 10°C and at 30°C. Radicles emerged from 86–100% of freshly matured seeds in light at 15, 20, 20/10 and 25/15°C within 28 days, but failed to emerge at 10°C. Radicles emerged equally well in a 12 h photoperiod and in continuous darkness at 25/15°C. Rapid embryo growth and germination over a range of conditions indicate that seeds of this taxon have morphological dormancy (MD); this is the first report of MD in a species of Lonicera . Seeds are dispersed in summer, at which time high temperatures promote embryo growth. Embryos grow to the critical length for germination in approximately 1 month; the peak of seedling emergence occurs in early autumn. Radicles emerged within 2 months from 98% of seeds buried at soil depths of 2 cm and 10 cm in the field in August in Sapporo, Japan; thus, seeds have no potential to form a persistent soil seed bank. However, seeds sown too late in autumn for embryos to grow remained viable and germinated the following summer when temperatures were high enough to promote embryo growth.     

Venation Pattern and Fruit Development in Lomatium dasycarpum (Umbelliferae)

Venation patterns of flower and fruit, and their relationshipto fruit development, are described for Lomatium dasycarpum(T. and G.) Coult. and Rose, a member of the tribe Peucedaneae(Umbelliferae). Comparisons of these observations are made withreports of fruit anatomy for other Peucedaneae, such as Angelica,Peucedanum, and Heracleum. These comparisons indicate that thereare distinctly different patterns of development leading toa mature fruit with a similar dorsal flattening. The possiblesignificance of such observations, with regard to our understandingof generic relationships in the tribe, is discussed.