Dormancy and germination responses of halophyte seeds to the application of ethylene

Ethylene is invariably produced during seed germination but its role in regulating seed dormancy and germination is poorly understood. Seeds of 22 halophytic species having different life forms – salt secreting dicots, salt secreting monocots, stem succulents and leaf succulents were germinated in Petri dishes kept in a growth chamber set at 20/30 °C (night/day) temperature and a 12 hr light period. Sodium chloride and ethephon were added to the medium from the beginning of the experiment. Seed germination was recorded every other day for twenty days. Application of ethylene did not have any significant effect on releasing seeds from innate dormancy. However, it appeared to have a role in alleviation of salinity effects which varied from negative in certain species to almost complete alleviation of high salinity effects in others.Our data indicates that ethylene appears to have little role in breaking innate seed dormancy however, in most halophyte seeds studied, application of ethylene alleviate the salinity effect to various degrees. Halophyte seeds which could germinate under saline conditions approaching twice the salinity of seawater may offer clues to understand management of seed germination under highly saline conditions. To cite this article: M.A. Khan et al., C. R. Biologies 332 (2009).     

The comparative germination ecology of nine Rumex species

The germination requirements, dormancy cycle and longevity of nine Rumexspecies were studied in field conditions and laboratory experiments to show theadaptations of the related species to their specific habitat. Within one genus,rather striking differences were observed in germination ecology. However, theclosely related species, R. acetosa and R.scutatus, are very similar: they fruit in early summer; theirseeds can germinate immediately after dispersal, and they are nondormant andshort-lived. R. acetosella also has fruits insummer, but the seeds do not germinate the first season after dispersal. Theyare long-lived, but buried seeds do not show a dormancy cycle; they mightgerminate in different seasons after exposure to light. Seeds of four species (R. conglomeratus,R. maritimus, R. sanguineus andR. crispus) are long-lived and undergo aseasonal dormancy cycle, with a low level of dormancy in winter and early springand a deep dormancy in summer as was already known for R.obtusifolius. These seeds are shed in the autumn, and they germinatemainly in the spring in consecutive years. R. maritimusalso germinates in summer and autumn on drying muddy soils. The seeds of R. hydrolapathum only germinate onwaterlogged soils, which explains its growth at the edge of streams and ponds.Its seeds are rather short-lived. The seeds of the species on very wetplaces require a higher temperature for germination.     

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.     

The importance of grazing intensity and frequency for physiological responses of the tropical seagrass Thalassia hemprichii

Seagrass grazing is an intrinsic disturbance in primarily tropical and subtropical areas. While there is a general parabolic response in seagrass growth to grazing intensity, there is less knowledge on the role of grazing frequency, as well as potential interactions between grazing intensity and frequency. This study experimentally investigated physiological responses in Thalassia hemprichii to simulated (leaf cutting) grazing regimes with different intensities (25% vs. 75%) and frequencies (1 times vs. 3 times) over 35 days in Chwaka Bay (Zanzibar, Tanzania). The results showed that the two high-intensity treatments (75% removal) had 37–41% lower growth rate than the low-intensity/low-frequency treatment, and rhizome sugar and starch content were both affected in a similar way. A 36% lower starch content in the simulated low-intensity/high-frequency regime (25% × 3) compared to the one of low-intensity/low-frequency (25% × 1) also shows an interaction between grazing intensity and frequency. This suggests that high-intensity (and to some extent frequency) grazing regimes, in comparison to low-intensity regimes, could negatively affect T. hemprichii growth, energy reserves, and thereby the ability to deal with additional stress like light limitation or grazing.     

The effect of cold stratification and light on the seed germination of temperate sedges (Carex) from various habitats and implications for regenerative strategies

The germination responses of 32 temperate Carex species were tested in light and darkness at five constant temperatures and under one fluctuating temperature regime, before and after cold-wet stratification. Using a linear logistic regression model, the probability of germination tested across all species was found to be significantly higher after stratification, in light and at the fluctuating temperature. In addition, the probability increased with temperature. Stratification increased germination in 28 species and had very little or no effect on four species. There was almost no germination in darkness prior to stratification, and the germination in light was considerably higher in all but two species compared with that in darkness. Thus, it can be concluded that the Carex species tested have broadly similar germination response patterns. The fact that Carices can be released from high levels of primary dormancy by low-temperature stratification implies that they are spring germinators. A light requirement after stratification in the major fraction of seeds and the capability of almost all investigated sedges to respond to fluctuating temperatures make it likely that persistent seed banks are formed. Additionally, sedges generally seem to have a high temperature requirement for germination which prevents them from emerging at the very beginning of the growing season. Regeneration by seed is probably largely restricted to gaps resulting from late spring disturbances where buried seeds have an opportunity to germinate and grow. Differences in germination were apparent between species occupying different habitats. Overall germination was significantly higher in wetland species than in dry-site species, probably owing to the greater capability of wetland species to respond to fluctuating temperatures. Differences in germination between forest and open-site species can be attributed to the higher capability of forest sedges to respond to low temperatures and temperature fluctuations. The influence of seed weight on germination was not significant in the 18 species adapted to wet, open habitats. There was, however, a tendency for the germination percentages to be low for large-seeded Carices. The interpretation of habitat differences is difficult due to a positive correlation between seed weight and dry habitats.     

Effects of seed storage on germination of two succulent desert halophytes with little dormancy and transient seed bank

Seeds of both Salsola imbricata and Haloxylon salicornicum have high germination level and germination speed, and form a transient seed bank in nature. The impacts of storage period and condition on germination level and speed were assessed in the two species. Storage for three months significantly increased both germination level and speed of seeds stored under the different conditions, compared to that of fresh seeds. In both species, nine months storage did not affect germination percentage in cold storage seeds, but completely inhibited it in field seeds. Storage for longer time in room and warm temperatures resulted in significant reduction or complete inhibition in the germination of the two species, so this was more pronounced in H. salicornicum. Storage significantly increased germinate rate index of seeds stored in all conditions till 17 months in S. imbricata and till 12 months in H. salicornicum. In both species, fridge storage had little effects on final germination and germination speed of seeds incubated at the different temperatures, compared to fresh seeds. However, room temperature and warm storages significantly reduced final germination and germination speed at the different temperatures, so the reduction was more pronounced at 35 °C, especially in H. salicornicum.     

Ecological genetics of seed germination regulation in Bromus tectorum L.

Regulation of seed germination phenology is an important aspect of the life history strategy of invading annual plant species. In the obligately selfing winter annual grass Bromus tectorum, seeds are at least conditionally dormant at dispersal in early summer and lose dormancy through dry-afterripening. Patterns of germination response at dispersal vary among populations and sometimes across years within populations. To assess the relative contribution of genotype and maturation environment to this variation, we grew progeny of ten parental lines from each of six contrasting populations in a common greenhouse environment. We then tested the germination responses of recently harvested seeds of the putative full-sib progeny at five incubation temperatures. Significant germination response differences among populations were observed in greenhouse cultivation, and major differences among full-sib families were evident for some populations and traits. Among-population variation accounted for over 90% of the variance in each trait, while within-family variance accounted for 1% or less. Germination responses of greenhouse-grown progeny were positively correlated with the responses of wild-collected seeds, but there was a tendency for lowered dormancy at higher incubation temperatures. This tendency was more marked in populations from cold desert, foothill, and plains habitats, suggesting a genotype-maturation environment interaction. Differences among populations in the amount of among-family variance were more evident at lower incubation temperatures, while among-family variance was more uniformly low at summer incubation temperatures. Populations from predictable extreme environments (subalpine meadow and warm desert margin) showed significantly less variation among families than populations from less predictable cold desert, foothill, and plains environments. Low among-family variance was not specifically associated with small population size or marginality of habitat, as small marginal populations from unpredictable environments showed variance as high as that of large populations. In populations with high among-family variance for germination traits, germination responses tended to be correlated across incubation temperatures, making it possible to characterize families in terms of their general dormancy status. The results indicate that seed germination regulation in this species is probably under strong genetic control, and that habitats with temporally varying selection are occupied by populations that tend to be more polymorphic in terms of their germination response patterns. Received: 19 May 1998 / Accepted: 27 January 1999     

Effects of pre-treatments and temperature on seed viability and germination of Juniperus macrocarpa Sm.

The effects of collecting season, collection site, laboratory pre-treatments and temperatures on seed viability and germination of Juniperus macrocarpa were investigated. Ripe cones were collected in four Sardinian dune systems, in two seasons, from plant and soil. Warm (W) and cold (C) stratifications, two combinations of them (W + C, C + W), and no pre-treatment (0) were applied. Seeds were incubated in a range of constant (10–25 °C) and an alternating (25/10 °C) temperature regime. Seed viability was low (ca. 40%) and varied significantly according to the collecting season. Seed germination was also low (ca. 10%), the 0 and W were the most effective pre-treatments on stimulating germination. The best germination temperature, without any pre-treatment, was 15 °C (ca. 20%). J. macrocarpa seeds are dormant and the achieved results suggested that the presence of secondary dormancy is induced by cold stratification. Spring appeared to be the best season for seed collecting, whereas autumn was the best for sowing. These results give new findings for restoration activities on this species.     

Tolerance and recovery responses of playa halophytes to light,salinity and temperature stresses during seed germination

Halogeton glomeratus (M. Bieb.) C.A. Mey., Lepidium latifolium Linn. and Peganum harmala Linn. are distributed in temperate salt playa habitats of Upper Hunza, Pakistan. Seeds were germinated under various salinity (0–500 mM NaCl), light (12 h-light:12 h-dark and 24 h-dark) and temperature (5/15, 10/20, 15/25, 20/30, and 25/35 °C, dark/light) regimes for 20 days to determine the optimal conditions for germination and recovery of seeds from these factors when exposed to less than optimal conditions. Seeds that failed to germinate in dark were transferred successively to 12 h-photoperiod, salinity to distilled water and from various temperature regimes to 20/30 °C, to determine the effect of these stresses and the ability of these seeds to recover respectively. Highest seed germination (H. glomeratus and L. latifolium: 100%; P. harmala: 80%) was obtained in non-saline control at 20/30 °C in 12 h-photoperiod, however, increase in salinity progressively inhibited seed germination. Seed germination of H. glomeratus and P. harmala was substantially inhibited and that of L. latifolium was prevented in dark. Salinity and dark treatments have a synergistic effect in inhibiting seed germination of all species. No seed of any species germinated at 5/15 °C; germination was substantially inhibited at 25/35 °C both for H. glomeratus and P. harmala while L. latifolium failed to germinate at 25/35 °C. Rate of germination also decreased with an increase in salinity at all temperature regimes but this effect was minimal at optimal temperature regime of 20/30 °C. After successive elimination of light, salinity and temperature stresses, final seed germination was identical to respective controls. The results indicate that seeds of these temperate halophytes could endure environmental stresses without losing viability and germinate readily when these stresses are removed. Under the extremely variable conditions of the playa habitat these species are highly opportunistic exploiting the windows of opportunity available during spring or early summer.     

The dual role of temperature in the regulation of the seasonal changes in dormancy and germination of seeds of Polygonum persicaria L.

Summary The role of temperature in the regulation of seasonal changes in dormancy and germination was studied in seeds of Polygonum persicaria. Seeds were buried in the field and under controlled conditions. Portions of seeds were exhumed at regular intervals and germination was tested over a range of conditions. Seeds of P. persicaria exhibited a seasonal dormancy pattern that clearly showed the typical features of summer annuals, i.e. dormancy was relieved at low winter temperatures, the germination peak occurred in spring and dormancy was re-induced in summer. The expression of the dormancy pattern was influenced by the temperature at which germination was tested. At 30°C exhumed seeds germinated over a much longer period of the year than at 20° or 10°C. Nitrate added during the germination test occasionally stimulated germination. The seasonal changes in dormancy of buried seeds were regulated by the field temperature. Soil moisture and nitrate content did not influence the changes in dormancy. The fact that, on the one hand, field temperature determined the changes in dormancy and, on the other hand, germination itself was influenced by temperature, was used to describe the seasonal germination pattern of P. persicaria with a model. Germination of exhumed seeds in Petri dishes at field temperature was accurately described with this model. Germination in the field was restricted to the period where the range of temperatures over which germination could proceed (computed with the model) and field temperature overlapped.     

The ecology of the seed in Trapa natans var. Japonica in a eutrophic lake

The production, density and environment of buried seeds and the temperature response of germinating seeds in Trapa natans var. japonica Nakai were ascertained. The mechanism of high reproductivity, tolerance of the plant for eutrophication and the stability of the stand in an eutrophic lake are discussed. The mean fresh weight of seeds was about 7.2 g, and the density of liver and dead seeds in April ranged 66.7 to 80.0 m^-2 and 26.7 to 97.8 m^-2, respectively. The threshold temperature that breaks dormancy and promotes germination in the lake was the low temperature, 8°C. The dormancy of buried seeds was almost broken under the field condition in January. The rate of germination obtained by the field experiment, and the estimation from the density of buried seed and the number of seeds germinated at the stand, were 87% and 60%, respectively. It was estimated that about 30% of the seedlings died before the floating leaf reached water surface. A rosette produced less than three seeds a year, which were fewer than for another Trapa, but the size of the former was larger. The number of seeds changed year by year, but the size-distribution pattern of seeds did not change with years, nor was it affected by the density of rosettes. The reason why T. natans var. japonica could succeed in eutrophic waters was attributed to reproductive strategy. The stand could recover by increasing the number of rosettes through the great branching ability of the main stem, even when the density of buried seeds was greatly decreased in the previous year.     

The control of seed germination in Trollius ledebouri: The breaking of dormancy

Seeds of Trollius ledebouri exhibit low germination when maintained on moistened filter paper. Dormancy can be overcome by the application of gibberellins A₄+A₇ or by testa removal. Germination is characterised by a change in the anatomy of the seed and by specific alterations in the protein complement of the endosperm tissue. These anatomical and biochemical changes are also exhibited by isolated endosperm tissue maintained in the absence of the embryo. The observations described are discussed in relation to the interaction between the endosperm and the embryo in the control of seed germination in T. ledebouri.Abbreviation GA gibberellin     

Morphology and Germination of Yellow and Brown Caryopses of Aegilops Geniculata Roth (Gramineae) Population from Italy

Abstract

The germination percentage of Aegilops geniculata Roth caryopses was studied on four Italian populations (Pisa, Manfredonia, Catania and Simbirizzi). Observation of the germination trend was carried out at constant temperatures of 10°, 20° and 30°C, from maturation up to 60 days after harvesting. Spike, spikelet and caryopsis phenotypic characteristics were also studied. The populations of Pisa, Manfredonia and Simbirizzi presented two different types of caryopses: yellow caryopses - larger, heavier and composing roughly 60% of total—and brown caryopses—smaller and lighter in weight. The Catania population formed an exception in that brown caryopses were virtually absent (5%). This Sicilian accession also showed the largest spikelet size.

Yellow caryopses germinated more rapidly than the dark ones, which were shown to be endowed with longer relative dormancy, above all at 10° and 30° C. Such a phenomenon allows the two types of caryopses to have a different germination ecology, thus providing A. geniculata with a broader choice of germination opportunities and consequently favouring spread and survival of the species.     

Modulation of germination of embryos isolated from dormant and nondormant barley grains by manipulation of endogenous abscisic acid

Dormant and non-dormant barley (Hordeum distichum L.) grains with identical genetic backgrounds were obtained by maturing grains under different climate conditions. When isolated embryos from dormant grains were incubated in a well containing a fixed volume of water (300 l), the germination rate and percentage were dependent on the embryo number per well. A higher embryo number per well was correlated with a lower germination rate and percentage. However, this was not the case for the embryos isolated from nondormant grains. During germination, the endogenous cis-abscisic acid (ABA) in isolated embryos from both dormant and nondormant grains was analyzed. The inhibitory effect on germination of a higher number per well of isolated dormant embryos was due to diffusion of endogenous ABA out of the embryos and accumulation of ABA in the incubation medium. Moreover, there was de-novo synthesis of ABA in embryos isolated from dormant grains during incubation but not in embryos isolated from nondormant grains. The inhibitory effect of ABA on germination of embryos isolated from dormant grains could be mimicked by addition of ABA or the medium in which dormant embryos had been placed. Embryos isolated from nondormant grains were insensitive to addition of ABA and medium from dormant embryos. Our results demonstrate that diffusion of endogenous ABA, de-novo ABA synthesis and ABA sensitivity play a role in the control of germination. It is proposed that dormancy-breaking treatments act via changes to these processes.Abbreviations ABA cis-abscisic acid - E/W embryo(s) per well Prof. K.R. Libbenga (Institute of Molecular Plant Sciences, Leiden University) is thanked for fruitful discussions. B.V.D. was partly supported by E.E.C. BIOTECH program PL 920175.     

The control of seed germination in Trollius ledebouri A model of seed dormancy

Treatment of Trollius ledebouri seeds with gibberellins A₄+A₇ promotes germination. The efficacy of the treatment is dependent upon the duration of imbibition in distilled water prior to GA₄₊₇ application. Presoaking increases both the final percentage germination attained and also its rate of achievement. No presoaking effect is exhibited by seeds induced to germinate by testa removal in the absence of GA₄₊₇. Active washing of Trollius seeds enhances the presoaking effect and the eluent from washed seeds is inhibitory to germination. The results support the hypothesis that the presoaking effect exhibited by Trollius is the result of the leaching of a germination inhibitor from the seeds which is antagonistic to GA₄₊₇. Additionally, treatment of Trollius seeds with the gibberellin-biosynthesis inhibitor (2-chloroethyl)-trimethylammonium chloride (CCC) prior to testa removal retards germination. The inhibitory effect of CCC on germination is overcome by GA₄₊₇. Although CCC inhibits embryo growth during the presoaking of intact seeds, it does not affect the increased sensitivity of presoaked seeds to GA₄₊₇. Therefore, although endogenous gibberellins may be involved in the germination process, they do not contribute to the presoaking phenomenon. The expansion of isolated endosperm tissue is not affected by CCC. However, the chemical markedly inhibits endosperm expansion in intact seeds and implicates the embryo as both the site of production of the germination inhibitor and of gibberellin. These results are discussed in relation to previous studies and a model is presented to account for the characteristics of germination in Trollius.Abbreviations GA gibberellin - CCC (2-chloroethyl)-trimethylammonium chloride     

种子休眠与萌发的分子生物学的研究进展

休眠与萌发是植物种子对环境变化的适应特征,受许多基因调控和环境因子的影响.利用数量遗传学方法(如QTL分析)和突变等手段已对休眠和萌发特性进行了深入的遗传学研究.近些年来,随着分子生物学的快速发展,种子休眠和萌发研究已经深入到分子水平.分子生物学技术的运用,特别是基因表达、基因组测序和以双向凝胶电泳及质谱分析为技术基础的蛋白质组学分析,已成为研究种子休眠和萌发的新工具和新方向.本文主要就利用分子生物学方法研究种子休眠与萌发的进展给予简要综述.     

Environmental influences on the formation and germination of hibernacula in the brackish-water bryozoan Victorella pavida Saville Kent, 1870 (Ctenostomata: Victorellidae)

Brackish-water and fresh-water bryozoans produce asexually derived dormant propagules that allow survival of unfavourable conditions and provide a potential means of dispersal. The propagules of brackish-water ctenostome bryozoans are called hibernacula. We monitored the life-cycle of the brackish-water ctenostome Victorella pavida Saville Kent, 1870 in its natural habitat and investigated, in laboratory cultures, the influence of temperature and salinity on the production and germination of hibernacula and on subsequent colony growth. V. pavida is a protected species in the UK, where its only locality is at Swanpool lagoon, Falmouth. Colonies were collected from Swanpool monthly from January 2004 to January 2005. Hibernaculum germination appeared to be triggered by increased water temperature (c. 13 °C) in the lagoon in March and April. In culture, germination was triggered by transfer from 5 °C to 19 °C in a range of salinities; subsequent colony growth was affected by salinity, with strongest growth at 13, 18 and 36 psu, and reduced growth at 5 and 9 psu. At 3.5 psu, hibernacula germinated, but there was no further development. At 36 psu there was an initial lag in growth, but after 30 d the colonies were comparable with those kept at 18 psu. Hibernaculum formation by colonies occurred from June to October, with production increasing towards October. Hibernacula appear not to have long-term viability but merely to permit survival from one year to the next. The results suggest that any changes in the hydrographic regime at Swanpool could have significant consequences for the survival of V. pavida.     

The growth physics and water relations of red-light-induced germination in lettuce seeds

Using lettuce (Lactuca sativa L., cv. Grand Rapids) embryos in osmotica, we have demonstrated that when the growth rates of the embryonic axes of seeds treated with red (R) or far-red (FR) light are equalized, the axes of R-treated seeds develop a 3.4-bar decrease in water potential (paper No. III).As axial growth begins, reserve protein and phytin decrease rapidly, concomitant with increases in reducing sugars, -amino nitrogen, and inorganic and esterified soluble phosphates. However, no differences between the axes of R-and FR-treated seeds are found with respect to the changes in these compounds, indicating that these changes arise as a result of growth and are not under immediate phytochrome control. Little change in the total lipid content is found in either treatment. The axes of FR-treated seeds hydrolyze endogenous sucrose at a greater rate thant those of R-treated seeds. Axes of R-treated seeds accumulate K⁺ and Na⁺ to a greater extent than those of FR-treated seeds. When potassium salts are added to the incubation medium, R induces increased K⁺ uptake by the axis and greater medium acidification by the axis. Malate and other organic acids and acidic amino acids increase at equal rates in both treatments, indicating that inorganic anions may also be taken up to balance the ionic charges. The results are compatible with the assumption that changes in the osmotic and pressure potentials of the embryonic axes of R-treated seeds are the result of a phytochrome-stimulated proton pump which, in whole dormant seeds, would initiate water-potential changes allowing the embryos to overcome the mechanical restraint of the surrounding seed layers, resulting in germination.Abbreviations FR far-red light - PEG polyethylene glyeol 4000 - Pfr far-red-absorbing form of phytochrome - R red light III=Carpita et al. 1979     

种子感光的机理及影响种子感光性的因素

种子的萌发或休眠均取决于萌发时种子内所建立起来的Pfr含量和Pfr/(Pr Pfr)比值。种子内的Pfr水平受到诸多因素的影响。光中性种子在成熟时已存在适合萌发的Pfr水平;需光种子在不同程度地接受白光或红光照射后,方可达到适宜的Pfr水平;忌光种子萌发要求的Pfr水平较低,因此萌发需要较长时间的黑暗。种子的感光性不是绝对的,母株的生长条件、种子本身的成熟度、贮藏状况、光质、光流量、光周期、萌发温度、O2供应及某些化合物的处理等都可使种子的感光性发生改变。