Ethylene as a Component of the Emanations From Germinating Peanut Seeds and Its Effect on Dormant Virginia-type Seeds

The embryonic axes of Spanish-type peanut seeds that do not exhibit dormancy to any extent were found to produce ethylene during germination. Virginia-type peanut seeds of the extremely dormant variety NC-13 produced low levels of ethylene when imbibed but not germinating. Treatments that released dormancy of NC-13 peanut seeds resulted in increased ethylene production by the embryonic axis. The estimated internal concentration of ethylene in Virginia-type peanut seeds was 0.4 ppm at 24 hr of germination. Fumigation with an external concentration of 3.0 to 3.5 ppm for 6 hr was sufficient to break dormancy of Virginia-type peanut seeds. These results suggest that ethylene is associated with the germination processes of non-dormant seeds and participates in the breaking of seed dormancy of dormant peanut varieties.     

Dormancy and enzyme levels in seeds of wild oats

Nine enzymes were compared in dry and steeped mature dormant and non-dormant seeds of wild oats. In dry seeds only glutamate-pyruvate transaminase and phosphoglycerate kinase were greater in non-dormant seeds. In steeped non-dormant seeds glucose-6-phosphate dehydrogenase activity doubled while the enzyme declined sharply in dormant seeds. Increases in isocitrate dehydrogenase, glutamate-oxaloacetate transaminase and acid phosphatase in non-dormant seeds, during steeping, are consistent with the hypothesis that the pentose phosphate and glycolysis-tricarboxylic acid pathways are involved in the control of dormancy of wild oat seed.     

Fructose 2,6-bisphosphate in germinating oat seeds. A biochemical study of seed dormancy

When dormant oat seeds were imbibed at the non-permissive temperature of 30 degrees C, the concentration of phosphoenolpyruvate and of glycerate 3-phosphate, which are two inhibitors of phosphofructokinase 2, increased almost linearly during 30 h. By contrast, these metabolites increased only after a lag period of about 10 h in non-dormant seeds imbibed at the same temperature. As a consequence of this, the concentration of the C3 derivatives remained always remarkably lower in non-dormant than in dormant seeds. Accordingly, the concentration of fructose 2,6-bisphosphate, which increased similarly in the two types of seeds during the first 8 h after the start of inhibition, then reached a plateau in dormant seeds but continued to increase for another 8 h in non-dormant seeds, reaching a maximal value a few hours before the beginning of radicle protrusion. When the dormant seeds were imbibed at the permissive temperature of 10 degrees C, the evolution of all metabolites was slowed down but behaved like that of non-dormant seeds imbibed at 30 degrees C. Experiments in which the dormant seeds were submitted to a jump from 10 degrees C to 30 degrees C and vice versa, always provoked reverse changes in the concentration of the C3 derivatives and of fructose 2,6-bisphosphate, the latter being increased in all conditions that allowed germination. Dormant seeds were also allowed to germinate at 30 degrees C by imbibition during 24 h in the presence of 3% ethanol. Again, this permissive treatment caused an arrest in the accumulation of C3 derivatives and an increase in fructose 2,6-bisphosphate. Another, apparently unrelated, biochemical difference between dormant and non-dormant oat seeds was their inorganic pyrophosphate content, which was approximately five-fold higher in non-dormant than in dormant seeds. This difference was observed before and persisted during imbibition as long as measurement could be made and was not affected by the temperature jumps or by ethanol. In contrast to the phosphoric esters under investigation, pyrophosphate was not preferentially located in the embryo.     

Role of reactive oxygen species in the regulation of Arabidopsis seed dormancy

Freshly harvested seeds of Arabidopsis thaliana, Columbia (Col) accession were dormant when imbibed at 25°C in the dark. Their dormancy was alleviated by continuous light during imbibition or by 5 weeks of storage at 20°C (after-ripening). We investigated the possible role of reactive oxygen species (ROS) in the regulation of Col seed dormancy. After 24 h of imbibition at 25°C, non-dormant seeds produced more ROS than dormant seeds, and their catalase activity was lower. In situ ROS localization revealed that germination was associated with an accumulation of superoxide and hydrogen peroxide in the radicle. ROS production was temporally and spatially regulated: ROS were first localized within the cytoplasm upon imbibition of non-dormant seeds, then in the nucleus and finally in the cell wall, which suggests that ROS play different roles during germination. Imbibition of dormant and non-dormant seeds in the presence of ROS scavengers or donors, which inhibited or stimulated germination, respectively, confirmed the role of ROS in germination. Freshly harvested seeds of the mutants defective in catalase (cat2-1) and vitamin E (vte1-1) did not display dormancy; however, seeds of the NADPH oxidase mutants (rbohD) were deeply dormant. Expression of a set of genes related to dormancy upon imbibition in the cat2-1 and vet1-1 seeds revealed that their non-dormant phenotype was probably not related to ABA or gibberellin metabolism, but suggested that ROS could trigger germination through gibberellin signaling activation.     

A meta-analysis of the effects of frugivory (endozoochory) on seed germination: role of seed size and kind of dormancy

Heretofore, no study has determined how germination of ingested seeds is affected by the kind (class) of dormancy nor by seed dormancy x seed size interaction. Thus, we aimed to determine the effects of seed size, kind of dormancy and their interaction on germination of defecated seeds using a meta-analysis. We collected data for 366 plant species in 97 plant families from 76 publications. In general, gut passage significantly increased germination percentage of defecated seeds by 5% compared with that of control seeds. Germination percentages of non-dormant, physiologically dormant, and morphologically/morphophysiologically dormant seeds (all water-permeable) significantly decreased after gut passage by 40, 18, and 14%, respectively, compared with control seeds (non-gut-passed). Changes in germination percentage of seeds with physical dormancy (water-impermeable) were positive, and gut passage increased germination by 69% compared with control seeds. Germination of small seeds decreased 8% after gut passage, whereas germination of both medium and large seeds increased by 18%. However, changes in germination percentage differed between categories of seed size in each class of dormancy. In physically dormant seeds, germination of all seed sizes improved after gut passage, and the magnitude of increase was higher for large than for medium and small seeds. Thus, gut passage increased germination of medium-size water-permeable seeds (physiologically dormant and morphologically/morphophysiologically dormant) more than it did for large and small seeds. However, gut-passage decreased or did not change the germination percentage of non-dormant seeds. Seed size and kind of dormancy should be included in studies on the effect of gut passage on germination.     

The influence of cytokinins on apple embryo photosensitivity and acid phosphatase activity during stratification

Kinetin (KIN) and benzyladenine (BA) stimulate to different extent the germination of apple embryos isolated from dormant seeds or seeds submitted to stratification. KIN is much more active in the replacement of light requirement in apple embryos germination. Both cytokinins decrease the photosensitivity of embryos isolated from the seeds stratified less than one month, but only BA accelerates the appearance of the second photosensitivity maximum, normally occuring on the 70th day of stratification. Both cytokinins stimulate the activity of acid phosphatase between the 30th and 50th day of apple seed stratification. The stimulation between the 50th day and the end of stratification is exerted only by KIN. These differences allow to discuss the specificity of action of particular cytokinins during the after-ripening and germination of apple embryos.     

Possible involvement of volatile compounds in the after-ripening of cocklebur seeds

The mechanism of emergence from primary dormancy, the process of after-ripening, in cocklebur (Xanthium pennsylvanicum) seeds was examined in relation to the involvement of volatile compounds and to the relative humidity (RH) in which the seeds were stored. The after-ripening of these seeds proceeds only at water contents between 7 and 14% which are conditioned under RHs of 33% to 53% and are identified with water-binding region II. After-ripening of cocklebur seeds occurred even in water-binding region I. imposed by 12% RH. when exposed to HCN gas during the storage period. Exposure of dormant seeds to acetaldehyde (ethanal) retarded after-ripening. even in water-binding region II. thus decreasing germinability. This decrease of germinability by ethanal was found also in the after-ripened seeds, suggesting that ethanal accelerates seed deterioration rather than retarding the after-ripening. The contents of ethanal. ethanal and HCN were high only in the dormant seeds held at 12% RH. Regardless of RH. a possible conversion of ethanal to ethanol. perhaps via alcohol dehydrogenase. was far larger in dormant than in non-dormant seeds. In contrast, the reverse conversion of ethanol to ethanal was more profound in non-dormant seeds. Pre-exposure of both types of seeds to HCN reduced the contents of both ethanal and ethanol at 12% RH. The contents of various adenylales including ATP in seed tissues were higher in dormant seeds stored at 12% RH than in non-dormant seeds after-ripened at 44% RH. It is suggested that emergence of cocklebur seeds from primary dormancy by HCN treatment at 12% RH may result from the reduction in the contents of ethanal via an unknown mechanism incurring the consumption of ATP. This implies involvement of volatile compound metabolism at the water-binding region II in the after-ripening process of cocklebur seeds.     

Ethylene in seed dormancy and germination

The role of ethylene in the release of primary and secondary dormancy and the germination of non-dormant seeds under normal and stressed conditions is considered. In many species, exogenous ethylene, or ethephon – an ethylene-releasing compound - stimulates seed germination that may be inhibited because of embryo or coat dormancy, adverse environmental conditions or inhibitors (e.g. abscisic acid, jasmonate). Ethylene can either act alone, or synergistically or additively with other factors. The immediate precursor of ethylene biosynthesis, 1-aminocyclopropane-1-carboxylic acid (ACC), may also improve seed germination, but usually less effectively. Dormant or non-dormant inhibited seeds have a lower ethylene production ability, and ACC and ACC oxidase activity than non-dormant, uninhibited seeds. Aminoethoxyvinyl-glycine (AVG) partially or markedly inhibits ethylene biosynthesis in dormant or non-dormant seeds, but does not affect seed germination. Ethylene binding is required in seeds of many species for dormancy release or germination under optimal or adverse conditions. There are examples where induction of seed germination by some stimulators requires ethylene action. However, the mechanism of ethylene action is almost unknown.
The evidence presented here shows that ethylene performs a relatively vital role in dormancy release and seed germination of most plant species studied.     

Physical forces in dormancy and germination of xanthium seeds

The germination of seeds of Xanthium pensylvanicum Wallr. occurs in 2 phases, an initial passive phase of water uptake followed by an active phase of growth. These 2 phases have been separated experimentally, and shown to occur similarly in isolated cotyledons and embryonic axes. Measurements of the physical thrust generated by the entire seed and its separate components of cotyledon and axis reveal that non-dormant Xanthium seeds develop more than twice the thrust of dormant seeds, and that this difference develops principally in the second phase of enlargement of the axis. Measurement of the forces required for piercing the testa of these seeds establishes that whereas the thrust developed by non-dormant seed is adequate to cause testa rupture, that developed by dormant seeds is not. It is concluded that the dormancy of Xanthium involves an inadequacy in the embryo for rupture of the testa.     

Dormancy in Cereal Seeds: II. THE NATURE OF THE GASEOUS EXCHANGE IN IMBIBED BARLEY AND RICE SEEDS

When barley seeds imbibe water, the O₂ uptake of non-dormantseeds is considerably less than that of dormant seeds for atleast the first 6 h, irrespective of the rate at which the seedshad previously lost dormancy. During the initial 6 h of imbibition, the CO₂ output of dormantbarley seeds is usually only slightly greater than and sometimesno different from that of nondormant seeds. The CO₂ output ofdormant seeds is reduced by about 66 percent by millimolar KCN,whereas that of non-dormant seeds is decreased by about 12–13per cent only. The CO₂ output of dormant barley in nitrogenis considerably less than the CO₂ output of non-dormant seedsunder the same conditions. Dormant rice seeds also show a higher initial O₂ uptake thannon-dormant seeds, though this is not generally as marked asin barley. Similarly, the initial CO₂ output of dormant seedsis distinctly greater than that of non-dormant seeds, but inmillimolar KCN it is depressed to a greater extent than in non-dormantseeds. In nitrogen, the CO₂ outputs of dormant and non-dormantseeds were found to be the same. Consequently, unlike barley,dormant rice seeds appear to be as capable of carrying out alcoholicfermentation under anaerobic conditions as nondormant seeds. In barley, increasing the O₂ tension from 21 per cent to 100per cent increased the oxygen uptake of dormant seeds more thanthat of non-dormant seeds (an increase of 53 per cent as against20–23 Per cent). In dormant seeds there was a concomitantincrease in CO₂ output (about 50 per cent), but the CO₂ outputof non-dormant seeds was hardly affected. High concentrations of CO₂ are inhibitory to the germinationof both dormant and non-dormant barley seeds. At a concentrationof 10 per cent, however, CO₂ is inhibitory only to dormant seeds,although at 2.5–5 per cent it is sometimes stimulatoryto the germination of dormant seeds. A 24–h treatmentwith appropriate concentrations of ethanol, lactic acid, oracetaldehyde is also stimulatory to the germination of dormantbarley seeds. Histochemical investigations in barley indicated the presenceof peroxidase, cytochrome oxidase, and -glycero-phosphate dehydrogenasein the embryo, aleurone layer, and in a layer associated withthe testa. A number of other redox enzymes were detected inthe embryo and aleurone layer only. No differences in distributionor intensity of activity were detected between dormant and nondormantseeds.     

Factors affecting the induction and release of secondary dormancy in Kalanchoë seeds

Abstract. Several short daily R irradiations are required from the first day of incubation on water to induce germination of Kalanchoë seeds. When the same light treatment is given after a prolonged dark incubation period at 20°C, secondary dormancy prevents germination. Factors controlling the induction and breaking of secondary dormancy have been investigated. The induction of secondary dormancy is very temperature dependent. Locally puncturing the seed coat strongly delays it. Secondary dormancy is not induced in the presence of GA₃ during the first 10 d of dark incubation, although this growth substance cannot induce dark germination. Prolonged or cyclic daily R irradiations can relieve secondary dormancy of seeds kept on water, even after a dark period of 20 d. A 24 h treatment at 4°C restores responsiveness to short R exposures of slightly secondarily dormant seeds. The synergism between GA₃ and Pfr in non-dormant Kalanchoë seeds, leading to high effectiveness of even one short FR irradiation, still occurs in seeds made secondarily dormant before transfer to GA₃, but more R or FR irradiations, in combination with GA₃, are required for the release of secondary dormancy. A combination of red light and 6-benzyl-aminopurine is ineffective in removing dormancy.     

Involvement of phytohormones in germination of dormant and non-dormant oat (Avena sativa L.) seeds

Oat seeds are susceptible to high temperature dormancy. Dormant grainsdo not germinate at 30 °C unless afterripened, dry, for severalweeks. Isolated embryos of dormant grains do germinate, especially ifGA₃ is added to the germination medium. ABA inhibits germinationproportionally to the concentration applied and GA₃ can overcome theABA inhibitory effect. Measurements of endogenous ABA and several GAs revealedthat the initial levels of ABA in dormant and non-dormant grains were quitesimilar. But, endogenous ABA in non-dormant seeds almost disappeared within thefirst 16 h of imbibition, while the amount in dormant grains haddecreased by less than 24%. The level of GA₁₉ in non-dormant seedswas higher, and GA₁₉ appears to be converted to GA₂₀ within the first 16h. The GA₂₀ was converted to GA₁ at leastduring the first 48 h of the germination process. Bothphytohormones thus appear to be involved in the germination process ofnon-dormant seeds. ABA first declines, while GA₁ is producedduring the first 16 h of imbibition to allow proper germination.Indormant grains the level of ABA remained high enough to prevent germinationduring at least a week and precursor GAs were not converted to GA₁.     

A Race for Survival: Can Bromus tectorum Seeds Escape Pyrenophora semeniperda-caused Mortality by Germinating Quickly?

BACKGROUND AND AIMS: Pathogen-seed interactions may involve a race for seed resources, so that seeds that germinate more quickly, mobilizing reserves, will be more likely to escape seed death than slow-germinating seeds. This race-for-survival hypothesis was tested for the North American seed pathogen Pyrenophora semeniperda on seeds of the annual grass Bromus tectorum, an invasive plant in North America. In this species, the seed germination rate varies as a function of dormancy status; dormant seeds germinate slowly if at all, whereas non-dormant seeds germinate quickly. METHODS: Three experimental approaches were utilized: (a) artificial inoculations of mature seeds that varied in primary dormancy status and wounding treatment; (b) naturally inoculated undispersed seeds that varied in primary dormancy status; and (c) naturally inoculated seeds from the carry-over seed bank that varied in degree of secondary dormancy, habitat of origin and seed age. KEY RESULTS: In all three approaches, seeds that germinated slowly were usually killed by the pathogen, whereas seeds that germinated quickly frequently escaped. Pyrenophora semeniperda reduced B. tectorum seed banks. Populations in drier habitats sustained 50 times more seed mortality than a population in a mesic habitat. Older carry-over seeds experienced 30 % more mortality than younger seeds. CONCLUSIONS: Given the dramatic levels of seed death and the ability of this pathogen to reduce seed carry-over, it is intriguing to consider whether P. semeniperda could be used to control B. tectorum through direct reduction of its seed bank.     

Redox-sensitive proteome and antioxidant strategies in wheat seed dormancy control

Oxidative signalling by ROS has been demonstrated to play a role in seed dormancy alleviation, but the detailed molecular mechanisms underlying this process remain largely unknown. Here, we show dynamic differences in redox-sensitive proteome upon wheat seed dormancy release. Using thiol-specific fluorescent labelling, solubility-based protein fractionation, 2-D IEF PAGE, and MS analysis in conjunction with wheat EST sequence libraries, proteins with reversible oxidoreductive changes were characterized. Altogether, 193 reactive Cys were found in 79 unique proteins responding differentially in dormant, non-dormant, abscisic, or gibberellic acid-treated seed protein extracts from RL4137, a wheat cultivar with extreme dormancy. The identified proteins included groups that are redox-, stress-, and pathogen-responsive, involved in protein synthesis and storage, are enzymes of carbohydrate metabolism, proteases, and those involved in transport and signal transduction. Two types of redox response could be detected: (i) a dramatic increase in protein thiol redox state in seeds during imbibition and hormonal treatment; (ii) higher antioxidant capacity related to sensing of a threshold redox potential and balancing the existing redox pools, in dry dormant versus non-dormant seeds. These results highlight occurrence of the antioxidant defence mechanisms required for the protection of seed during a dormancy stage.     

Herbaceous plant strategies in disturbed habitats

A systematic theoretical evaluation has been made of three important plant life history traits: adult longevity, seed longevity and seed mass, where seed mass is interpreted as being indicative of dispersal distance and seedling vigour. This model study examined the role of these three traits in relation to environmental disturbance. We chose temperate grasslands, widespread in north Western Europe and northern and eastern America, as our reference system for our simulations. Eight plant strategies were defined by allowing two levels in each of the three and combining them in all eight possible ways. A simple, spatially explicit model was developed to simulate competition among individuals with these eight trait combinations at different levels of disturbance.
Simulation results were compared with the actual occurrence over a disturbance gradient of species with similar plant trait combinations in a large database from the Sheffield area (UK). This showed that with increasing disturbance level, non-dormant perennials, dormant perennials, non-dormant annuals and dormant annuals, respectively, became dominant but only if small-seeded, indicating the relative viability of these particular strategies with respect to disturbance.
A new prediction from the model was that stable coexistence occurs between plant strategies with dormant and with non-dormant seeds over a range of levels of disturbance. Plant strategies with large seeds were inferior to small-seeded ones if competitive ability of seedlings is proportional to seed weight. This difference was highest at low seed densities and low germination probabilities, indicating that large-seeded species secure no advantage from being dormant (i.e. having a low germination probability). Finally, the results indicated that dormancy is superior to dispersal as a method of coping with disturbance.     

No contribution of the pentose phosphate pathway in dormancy-breaking of cocklebur seeds

The role of the oxidative pentose phosphate (PP) pathway in the dormancy-breaking of cocklebur (Xanthium pennsylvanicum Wallr.) seeds was investigated. D-[1-¹⁴C]-glucose or D-[6-¹⁴C]-glucose was fed to dormant and non-dormant lower seeds or to their axial or cotyledonary segments which were imbibed for different durations, and C₆/C₁ ratios of respired ¹⁴CO₂ as an index of the PP pathway activity were calculated. Contrary to expectation, there was no significant difference in the C₆/C₁ ratios between the dormant and non-dormant seeds or segments during a water imbition period of 24 h, although the PP pathway actually operated already in an early stage of water imbibition. Also concerning the activities of G6PDH and 6PGDH, the key enzymes of this pathway, no difference between the dormant and non-dormant seeds was found. It was thus concluded that, unlike other seeds, there is no contribution of the PP pathway to the regulation of dormancy of the cocklebur seed.     

Soil burial induced dormancy in weedy rice seeds through hormone level changes: Implications in adaptive evolution and weed control

Seed dormancy plays a key role in preventing seeds of higher plants from random germination under adverse environmental conditions. Previous studies suggested that a critical temperature could regulate germination of weedy rice seeds without primary dormancy at seed dispersion. However, what will happen to the non-dormant seeds after shattering in the soil seed banks when temperature fluctuates to exceed the critical temperature remains an interesting question to be investigated. To determine whether or not soil burial can change the status of dormancy in weedy rice seeds, we examined germination ratios of weedy rice seeds after soil-burial treatments. In addition, we compared hormone levels in the untreated seeds and viable but ungerminated seeds after soil burial. Results showed that soil burial induced a proportion of 41%–72% dormant seeds in the initially non-dormant weedy rice seeds. Also, the induction of seed dormancy is associated with the change of hormone levels in the seeds treated by soil burial, suggesting that soil burial can significantly activate the control of hormone production in seeds. Together, the previously reported mechanism of critical temperature-inhibited seed germination and the newly found phenomenon of soil burial-induced seed dormancy provide a “double-security” strategy to ensure germination of weedy rice seeds under a favorable condition in agricultural ecosystems. The findings not only reveal the important role of rapid evolution of adaptive functions in weeds, such as weedy rice, in adapting to changing agricultural environments, but also facilitate the design of strategies for effective weedy rice control practices.     

Glycolytic activity in embryos of Pisum sativum and of non-dormant or dormant seeds of Avena sativa L. expressed through activities of PFK and PPi-PFK

A quantitative cytochemical assay for PPi-PFK activity in the presence of Fru-2,6-P2 is described along with its application to determine levels of activity in embryos of Pisum sativum and Avena sativa. The activity of ATP-PFK has also been studied in parallel as have PFK activities during the switch from dormant to non-dormant embryos in Avena sativa. PPi-PFK activity has been demonstrated in all tissues of Pisum sativum embryos and of Avena sativa embryos including the scutellum and the aleurone layers. The PPi-PFK activity was greater than that of ATP-PFK in both dormant and non-dormant seeds though with only marginally more activity in the dormant as opposed to the non-dormant state.     

Studies on the Germination of Cereals 5. The Dormancy of Barley Grains during Ripening

The germinability of two barley varieties with different dormancycharacteristics has been determined during ripening, under fieldconditions on the plant, and under conditions of controlledhumidity after premature separation. In both cases, primarydormancy was lost more rapidly in the non-dormant variety (Domen)than in the dormant variety (Herta), and under conditions wherethere was an early reduction in the moisture content of thegrains. When the moisture content also fell to a lower levelthan in the field, primary dormancy in both varieties was almosteliminated by the normal time of harvest. Secondary dormancy,with recovery during dry storage, was induced in some grainsof the dormant variety, and also in a few of the non-dormantvariety, when their moisture content was increased by wet weatherat the harvest ripe stage. During the early stages of ripening, separation from the plantcaused an earlier reduction in the moisture content and an earlierincrease in the germinability of the grains of both varieties.The non-dormant variety, however, reached the same level ofgerminability, at the normal time of harvest, irrespective ofthe time of separation, whereas the dormant variety showed aprogressive decrease in germinability as the time of separationwas delayed. The condition of total primary dormancy during the initial growthof the grains was imposed by the covering layers, and therewas no varietal difference in the ability of the embryos togerminate when exposed. But when the moisture content of intactgrains was reduced at this stage, by separation from the plant,the same varietal difference in germinability became apparent.The possible effects of desiccation and the characteristicsof the two varieties are discussed in relation to the dormancyof grains when harvested.