Seed germination in Chenopodium album L.: further evidence for the dependence of the effects of growth regulators on nitrate availability

Abstract Chenopodium album L. plants, grown under controlled environmental conditions on different levels of soil nitrate, produced seeds with proportionately different NO^?₃ contents. Regardless of the endogenous NO^?₃ content, few seeds germinated in water or upon treatment with KNO₃. Ethylene promoted germination, and the extent of germination was positively correlated with the endogenous seed NO^?₃ content. Combined application of ethylene and KNO₃ in the dark had a synergistic effect on NO^?₃ -deficient seed. The synergism between ethylene and KNO₃ was attributable to the NO^?₃ moiety of the nitrate salt. Ethylene and light showed moderate synergism in seeds with low or high endogenous nitrate. Addition of nitrate, however, masked the interaction between ethylene and light. Gibberellic acid₄₊₇ (GA₄₊₇) or red light, each alone or combined with KNO₃, had little effect on germination. When applied together in the dark, ethylene and GA₄₊₇ synergistically enhanced the germination of NO^?₃-deficient seed. The combined effects of the two hormones on this seed were further enhanced by the addition of KNO₃. There was no synergism between ethylene and GA₄₊₇ in NO^?₃-rich seed. These interactions among GA₄₊₇, ethylene and KNO₃ were not affected by light. The results confirm and further elaborate our earlier finding that the sensitivity of C. album seeds to ethylene may depend on nitrate availability.     

Effects of salinity and nitrate on production and germination of dimorphic seeds applied both through the mother plant and exogenously during germination in Suaeda salsa

Salinity and nitrogen are two important environmental factors that affect the distribution of halophytes in their natural saline habitats. Seeds of the euhalophyte Suaeda salsa L. were harvested from plants that had been treated with 1 or 500 mm NaCl combined with 0.5 or 5 mm NO₃^?‐N (nitrate) for 115 days in a glasshouse. Germination was evaluated under different concentrations of NaCl and nitrate. Plants exposed to high salinity (500 mm ) and low nitrate (0.5 mm ) tended to produce heavy seeds. Either high salinity (500 mm ) or high nitrate (5 mm ) increased the brown/black seed ratio. The concentrations of Na⁺, K⁺, and Cl^? were higher in brown than in black seeds, and NO₃^? concentrations were higher in black than in brown seeds, regardless of NaCl and nitrate treatments during plant culture. Regardless of NaCl and nitrate concentrations during germination, seeds from plants grown with 0.5 mm nitrate generally germinated more rapidly than seeds from plants grown with 5 mm nitrate, and the difference was greater for black than for brown seeds. Exogenous nitrate during germination enhanced the germination of brown seeds less than that of black seeds. Producing more brown seeds and heavy black or brown seeds appears to be an adaptation of S. suaeda to saline environments. Producing more black seeds, which tend to remain dormant, should reduce competition for nitrogen and appears to be an adaptation to nitrogen‐limited environments. In conclusion, nitrate provided exogenously or by mother plants to black seeds may act as a signal molecule that enhances the germination of black S. suaeda seeds.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.     

THE EFFECT OF SEED DIMORPHISM ON THE GERMINATION AND SURVIVAL OF SALICORNIA EUROPAEA L. POPULATIONS

Seed dimorphism influenced the germination behavior of Salicornia europaea L., with small seeds being more dormant and less salt tolerant than large seeds. All of the large less dormant seeds of S. europaea germinated prior to May, and all seedlings produced after this time were from small seeds. A persistent seed bank was maintained by the small dormant seeds. Survivorship was relatively constant during the normal germination season, but increasing salinity stress at any time during the spring reduces the chance of seedling survival. Little germination occurred from July through September because of the high surface soil salinities during this period.     

Factors affecting the decomposition of 2-chloroethylphosphonic acid in soils

Abstract The ethylene-releasing compound 2-chloroethylphosphonic acid (ethephon) stimulates germination of dormant seeds under laboratory conditions and it may prove to be an effective means of administering ethylene to promote germination of seeds in the field. We examined the decomposition of ethephon in different soils to determine which properties of soils influence the activity (ethylene release) of ethephon. The release of ethylene from ethephon was not affected by the type of soil, the texture, the presence of organic matter, or levels of extractable SO^2?₄, PO^3?₄ NO^?₃, K⁺ or Na⁺. The rate of decomposition was, however, dependent on soil pH and was influenced by the soil moisture content. Very little ethylene was released from ethephon in soils with a pH <7.7 and with a moisture content < 10%.     

Seed germination and in vitro propagation of Maytenus canariensis through regeneration of adventitious shoots from axillary and apical buds

Seed germination and micropropagation protocols of the medicinal species Maytenus canariensis (Loes.) G. Kunkel & Sunding were optimized. In vitro seed germination occurred (86 to 94.7 %) only after treatment of the seeds with H₂SO₄, followed by surface sterilization and culture on solid nutrient medium without any growth regulators. Micropropagation failed when explants were taken from mature trees, and browning of the nutrient medium frequently occurred despite testing many growth media. Nonetheless, adventitious shoot regeneration was achieved employing axillary or apical buds taken from 2–2.5 months old plantlets obtained after in vitro germination of seeds, following culture on nutrient media supplemented with benzylaminopurine, kinetin and naphthalenacetic acid (NAA), attaining up to 3.9 shoots per explant, after 4–6 months. Root induction was best on a medium containing 4.0 mg dm⁻³ NAA, achieving a 100 % induction. After hardening of rooted plants, survival after transfer to soil was 71.43 %.     

Germination studies on three critically endangered endemic angiosperm species of the Kashmir Himalaya,India

The critically endangered and perennial alpine endemic angiosperms, namely, Aquilegia nivalis, Lagotis cashmeriana and Meconopsis latifolia inhabit such habitats in the Kashmir Himalaya that are characterised by short growing season and heavy snow cover for about 3–4 months during winter season. The seeds of these species under natural conditions experience a long period of pre-chilling during winter prior to their germination in following spring season. Taking cue from such a requirement, present study investigated the effect of chilling and exogenous application of growth hormones, NO₃ ⁻ and NH₄ ⁺ on total percent germination of otherwise deep-dormant seeds of these species, under alternate light/dark and continuous dark light regimes. Prolonged pre-chilling followed by treatment of seeds with different doses of GA₃ had a pronounced stimulatory effect on the total germination percentage in all the three species. In fact, highest germination percentage in A. nivalis was recorded only when pre-chilled seeds were treated with 1.5 mM GA₃ under alternate light/dark conditions. Likewise, germination of pre-chilled seeds of L. cashmeriana with ruptured seed coats was improved when treated with various concentrations of GA₃. Seed germination in M. latifolia was also favourably influenced by treatment of pre-chilled seeds with GA₃ or nitrogen applied either as NO₃ ⁻ or as NH₄ ⁺ under alternate light/dark conditions. Treatment of seeds with kinetin (6-furfuryl-aminopurine) had no significant influence on germination percentage in any of the three species. Thus, prolonged chilling of seeds followed by their treatment with GA₃ under alternate light/dark conditions are the requirements necessary for seed germination in these species.     

Root nitrogen concentration of sorghum above 2% produces least Striga hermonthica seed stimulation

A series of pot and laboratory experiments was carried out to assess the effects of N status of sorghum roots and timing of N application (as NH₄NO₃) on the germination of Striga hermonthica seeds. Root N concentrations varied from 10 to 26 mg N g^?1. The cut root and the root exudates technique used in assaying S. hermonthica seed germination gave similar results. However, the cut root technique was easier to handle and more discriminating at low germination levels. S. hermonthica seed germination per unit sorghum root mass followed a broken‐stick model. It decreased with increasing root N concentrations, reaching lowest levels at a root N concentration of 19.5 mg N g^?1, after which no further reduction occurred. It was not possible to reduce S. hermonthica seed germination to a zero level. Timing of N application influenced the time a higher N concentration is reached, not the S. hermonthica seed germination. Both timing and rate of N application are important in maintaining root N concentrations above 19.5 mg N g^?1, thereby potentially reducing S. hermonthica germination in the field. Translation of results to reductions in infection levels and yield losses is hampered by density‐dependent relations after the S. hermonthica germination stage.     

Effects of ethylene and some other environmental factors on different stages of germination in red root pigweed (Amaranthus retroflexus L.) seeds*

Abstract. Ethylene was found to promote two distinct processes during germination of redroot pigweed (Amarantus retroflexus L.) seeds: embryo expansion that splits the seed coat (incomplete germination), and radicle penetration through the more elastic endosperm (complete germination). The two events can be separated in time by subjecting seeds to low water potential or low CO₂ levels, which arrest germination of some seeds at the incomplete stage. Ethylene applications to incompletely germinated seeds promote complete germination, with a response threshold near 0.02 cm³ m^?3 and saturation near 0.5 cm³ m^?3. Higher ethylene concentrations (0.5 to 50 cm³ m^?3) given during the first day of seed imbibition also increase the percentage of seeds which initiate embryo expansion and split the seed coat. Light and elevated CO₂ also promote radicle penetration of the endosperm in seeds incubated under water stress. The results support the view that the germination pause at the incomplete stage is an adaptation to environmental stresses that can be overcome with exogenous ethylene or certain other stimuli.     

Seed yield,seed size and germination behaviour in the annualPogogyne abramsii

Summary Models of the evolution of seed dormancy reveal that dormancy is favoured either when opportunities for establishment vary over time and when there is wide variation in the probability of success, or when the probability of success is limited by frequency dependence. Empirical evidence supporting the temporal heterogeneity hypothesis exists, but there is scant evidence for dormancy being favoured by frequency dependent competition among seedlings. We test the hypothesis that the intensity of between-sib competition should favour a positive relationship between maternal fecundity and seed dormancy. This hypothesis is supported for the rare, vernal pool annual,Pogogyne abramsii: the proportion of dormant offspring was significantly higher among high fecundity mothers than among other mothers. Dormancy inP. abramsii is controlled by the seed coat, a maternal tissue, so delaying germination favours the inclusive fitness of mothers by reducing the potential for competition among siblings. Seed weight and time to first germination varied significantly amongP. abramsii plants and mean seed weight increased linearly with plant biomass. Seed weight and seed number are independently regulated by plant size. Overall, seed weight varied 10-fold and variability in seed weight within mothers was not explained by plant biomass, seed yield or mean seed weight. GerminableP. abramsii seeds were significantly heavier than dormant seeds, and germinable seeds heavier than 0.20 mg germinated more rapidly than those smaller than 0.20 mg.     

Seed viability and seed dormancy of non-native Phragmites australis in suburbanized and forested watersheds of the Chesapeake Bay, USA

The non-native, invasive haplotype of Phragmites australis is rapidly invading tidal and non-tidal wetlands across North America. Phragmites has the potential to spread by seeds and rhizomes. Seed viability and dormancy differences were quantified among 18 patches of non-native Phragmites in subestuarine wetlands in developed (i.e., suburbanized) vs. forested watersheds of the Chesapeake Bay. We used tetrazolium and germination assays to assess seed viability and compared germination percentages and rate of germination among fresh seeds, cold–moist treated seeds, and warm–dry treated seeds to evaluate seed dormancy. Seed viability was <1% in most patches but a few patches produced abundant viable seeds (5–21%). Seed viability, however, did not differ significantly between wetlands in forested vs. developed watersheds. Contrary to studies of Phragmites seed dormancy in European populations, some Phragmites seeds were dormant at maturity; cold–moist treated seeds germinated faster and to higher percentages than fresh seeds or warm–dry treated seeds.     

SEED DORMANCY AND GERMINATION IN MELAMPYRUM LINEARE

Immature seeds of Melampyrum lineare Desr. have very high germination percentages and dormancy is induced in a variable fraction of the seed crop during ripening. Correlated with this is the endogenous gibberellin-like activity which is found in considerable amounts in immature seeds, less in batches of ripe seeds, and is not detectable in batches containing only dormant seeds. For germination dormant seeds require activation followed by cold storage. In the laboratory activation is produced by allowing moist, dormant seeds to respire freely for several weeks at 20 C, or by treatment with exogenous GA₃. Dormancy appears to be most directly related to inability of the embryo to hydrolyze the thickened, mannan-containing endosperm cell walls. Embryos excised from dormant seed can be grown on agar enriched with whole macerated dormant seeds or with the ethanol-extractable materials from these (mostly sucrose and a glycoside). However, dormant seed material does not support growth when extracted to remove benzene- and ethanol-soluble materials.     

盐氮处理下盐地碱蓬种子成熟过程中的离子积累和种子萌发特性

研究了盐氮处理条件下盐地碱蓬种子成熟过程中的离子积累以及种子萌发特性,以理解盐地碱蓬在种子发育及萌发过程中对高盐低氮生境的适应性。结果表明,种子成熟过程中,不同浓度盐氮处理下(0.5和5 mmol/L NO₃^--N;1和500 mmol/L NaCl),与果皮和果枝相比, 胚中Na⁺、K⁺、Cl^- 和NO₃^-离子含量几乎没有变化。所有盐氮处理下Na⁺ 和Cl^-都是果皮和果枝中高于胚中,尤其是在高盐处理下。高盐处理下,K⁺ 和NO₃^-含量呈现相反的趋势。高氮时无论高盐还是低盐,果皮中NO₃^-离子含量高于胚中,而果枝中NO₃^-离子含量低于胚中。而低氮时果皮及果枝中NO₃^-离子含量均显著低于胚中。与高氮环境下收获的种子相比,低氮环境下收获的种子萌发率,萌发指数,活力指数都要明显高。上述结果说明,盐地碱蓬种子成熟过程中存在完善的离子调控机制,保护胚免受Na⁺ 和Cl^-等有害离子的伤害并且促进K⁺ 和NO₃^-等营养离子的积累。低NO₃^--N下收获的种子对外界的NO₃^-含量比较敏感,施以较高浓度的NO₃^-能够促进种子萌发,提高萌发指数和活力指数,可能与盐地碱蓬长期适应高盐低氮生境有关。     

The chemical nature of P accumulation in agricultural soils��implications for fertiliser management and design: an Australian perspective

Sand burial, persistent seed bank and soil water content (SWC) are three factors that potentially can affect regeneration in sand dune plant populations. To evaluate the effects of these three factors on population regeneration of Eremosparton songoricum, a rare and endangered legume, we investigated seed germination, seedling emergence and seedling survival in greenhouse and controlled field experiments in different sand dunes microsites. Freshly matured seeds are physically dormant, and the highest germination was only 9.3?±?5.8% at 25/10°C. Seed germination occurred at burial depths from 0 to 10 cm, but the maximum depth from which seedlings emerged was 6 cm; from 1 to 6 cm, the deeper the burial, the lower the percentage of seedling emergence. Only 36.2% of the total soil seed banks occurred at depths of 0?C6 cm. For artificially sown seeds at different dune microsites, mean seedling emergence percentage was 6.8%. Of 150 seedlings that emerged in the field investigation at the study site, only those germinating in flat sandy areas survived, and mean survival percentage was only 2.0%. Thus, the proportion of non-dormant seeds in soil seed banks that developed into seedlings and survived to the end of the growing season was only 0.2%. Minimum SWC for seed germination, seedling emergence and seedling survival was 2.0%. During monitoring of emergent seedlings in the field, low seedling recruitment was at least partly due to the rate of root extension (1.6?±?0.3 cm day^?1) into the sandy soil, which was slower than that of the downward movement of plant-available moisture (2.8?±?0.6 cm day^?1). Thus, population regeneration under natural conditions rarely occurred via sexual reproduction, owing to the limited water resources available for seedling establishment. Rational field seeding practices, including manually scarified or dry stored seeds before sowing, sowing the seeds at right time and suitable place, are suggested for ecological restoration of endangered E. songoricum populations.     

Seed development and germination ecophysiology of the invasive tree <Emphasis Type="Italic">Prunus serotina</Emphasis> (Rosaceae) in a temperate forest in Western Europe

Seed development, dormancy and germination of the American invasive tree species, Prunus serotina, are described for plants growing in a large forest in Belgium. Seeds of P. serotina were collected following anthesis in the first week of July and thereafter at fortnightly intervals. Seed dormancy, temperature requirements for germination and the soil seed bank were investigated. At maturation (about 105 days after anthesis), seed moisture content had decreased to around 13.7%, and 44% of the seeds had attained the capacity to germinate. Mature seeds of P. serotina exhibited physiological dormancy, germinating only after a long cold, moist stratification period. Highest germination percentage occurred in seeds treated with gibberellic acid (GA₃), at 10°C. We found no evidence that P. serotina forms a persistent seed bank but noticed a persistent seedling bank in the field.     

Natural regeneration of Spanish black pine [Pinus nigra Arn. ssp. salzmannii (Dunal) Franco] at contrasting altitudes in a Mediterranean mountain area

Information on plant seed dispersal, natural loss dynamics of seeds and germination are critical for understanding natural regeneration mechanisms. The aim of this study was to assess the effect of different forest stand densities on seedfall, seed predation, and seedling germination of two populations of the endangered Spanish black Pine forests located at lower (Central population) and higher elevation near the limit of the species?? range (peripheral population) in the Cuenca Mountains of Central Spain. The seed predation and germination experiment also included a nested site preparation treatment. Seed fall varied significantly between 2006 and 2005 or 2007 in both populations. During the only mast year of 2006, higher seedfall was observed at lower elevation and in higher density stands. Predation rates were influenced by the seed crop since predators consumed more than 75?% of seeds in years with lower production and less than 15?% in a mast year. Seed germination is influenced by forest habitat, stand density and soil scalping. For common habitat types, and in a high seed production year, better seed germination rates were observed in medium and dense stands (25?C30 and 35?C40?m²?ha^?1, respectively, in terms of basal area). No statistical difference in seed germination rate was found for Spanish black pine forest at its ecological distribution limit between lower and higher densities (15?C20 and 35?C40?m²?ha^?1, in terms of basal area). In both sites, closed stands with soil scalping exhibited higher germination rates.     

ABA, GA3, and nitrate may control seed germination of Crithmum maritimum (Apiaceae) under saline conditions

Impaired germination is common among halophyte seeds exposed to salt stress, partly resulting from the salt-induced reduction of the growth regulator contents in seeds. Thus, the understanding of hormonal regulation during the germination process is a main key: (i) to overcome the mechanisms by which NaCl-salinity inhibit germination; and (ii) to improve the germination of these species when challenged with NaCl. In the present investigation, the effects of ABA, GA₃, NO⁻₃, and NH⁺₄ on the germination of the oilseed halophyte Crithmum maritimum (Apiaceae) were assessed under NaCl-salinity (up to 200 mM NaCl). Seeds were collected from Tabarka rocky coasts (N-W of Tunisia). The exogenous application of GA₃, nitrate (either as NaNO₃ or KNO₃), and NH₄Cl enhanced germination under NaCl salinity. The beneficial impact of KNO₃ on germination upon seed exposure to NaCl salinity was rather due to NO⁻₃ than to K⁺, since KCl failed to significantly stimulate germination. Under optimal conditions for germination (0 mM NaCl), ABA inhibited germination over time in a dose dependent manner, but KNO₃ completely restored the germination parameters. Under NaCl salinity, the application of fluridone (FLU) an inhibitor of ABA biosynthesis, stimulated substantially seed germination. Taken together, our results point out that NO⁻₃ and GA₃ mitigate the NaCl-induced reduction of seed germination, and that NO⁻₃ counteracts the inhibitory effect of ABA on germination of C. maritimum. To cite this article: A. Atia et al., C. R. Biologies 332 (2009).     

Some effects of nitrate-treated soil upon the sensitivity of buried redroot pigweed (Amaranthus retroflexus L.) seeds to ethylene, temperature, light and carbon dioxide

Abstract Fresh dormant redroot pigweed (Amaranthus retroflexus L.) seeds were buried 5 cm deep in the field at Stoneville, MS in November 1981. Potassium nitrate (200 kg ha ¹) or nothing was applied to the soil in the fall of 1981 and the late winter of 1982. Seeds were recovered at intervals under darkness during the following 2 years and tested for responses to ethylene, temperature, light and carbon dioxide. During the first overwintering, nitrate enhanced loss of primary dormancy and increases seed sensitivity to temperature, light and ethylene. The loss of dormancy reached a maximum at 25 to 30 weeks (early summer) after burial. Examination of the recovered seeds indicated that about 80% of the non-treated seeds and 98% of the nitrate-treated seeds germinated in situ during the period of maximum loss of dormancy. Thus, after one overwintering period, about 20% of the original buried seed population remained dormant in nontreted soil and 2% remained dormant in the nitratetreated soil. After the second overwintering, the percentages of dormant seeds remaining in nontreated or treated soil were both only 1–2%. Nitrate reduced dormancy and enhanced germination in early summer following the first overwintering. Regardless of treatment, the remaining 1 2% of seeds in soil after the second year were of low sensitivity to the germination stimuli (ethylene, temperature, light) and constituted the long-lived portion of the original seed population.     

Response of Amaranthus retroflexus L. seeds to gibberellic acid, ethylene and abscisic acid depending on duration of stratification and burial

Freshly harvested, dormant seeds of Amaranthus retroflexus were unable to germinate at 25 and 35 °C. To release their dormancy at the above temperatures, the seeds were stratified at a constant temperature (4 °C) under laboratory conditions or at fluctuating temperatures in soil or by outdoor burial in soil. Fully dormant, or seeds stratified or buried (2006/2007 and 2007/2008) for various periods were treated with exogenous gibberellic acid (GA₃), ethephon and abscisic acid (ABA). Likewise, the effects of these regulators, applied during stratification, on seed germination were determined. The results indicate that A. retroflexus seed dormancy can be released either by stratification or by autumn–winter burial. The effect of GA₃ and ethylene, liberated from ethephon, applied after various periods of stratification or during stratification, depends on dormancy level. GA₃ did not affect or only slightly stimulated the germination of non-stratified, fully dormant seeds at 25 and 35 °C respectively. Ethylene increased germination at both temperatures. Seed response to GA₃ and ethylene at 25 °C was increased when dormancy was partially removed by stratification at constant or fluctuating temperatures or autumn–winter burial. The response to GA₃ and ethylene increased with increasing time of stratification. The presence of GA₃ and ethephon during stratification may stimulate germination at 35 °C. Thus, both GA₃ and ethylene can partially substitute the requirement for stratification or autumn–winter burial. Both hormones may also stimulate germination of secondary dormant seeds, exhumed in September. The response to ABA decreased in parallel with an increasing time of stratification and burial up to May 2007 or March 2008. Endogenous GA_n, ethylene and ABA may be involved in the control of dormancy state and germination of A. retroflexus. It is possible that releasing dormancy by stratification or partial burial is associated with changes in ABA/GA and ethylene balance and/or sensitivity to these hormones.