Characteristics of alternating temperatures which stimulate loss of dormancy in seeds of Rumex obtusifolius L. and Rumex crispus L.

Abstract. Alternating temperatures stimulate the germination of Rumex crispus L. and Rumex obtusifolius L. The optimum period spent at the lower temperature in a diurnal cycle is greater than that spent at the higher temperature. Under most conditions the optimum period at the upper temperatures is about 8 h but, as the upper temperature of a cycle is increased, the optimum period at the upper temperature becomes shorter and more critical. Thus when it is 35°C the optimum period is 2.5–4 h in the light, or about 1 h in the dark. The effect of alternating temperatures is much less in the dark than in the light and in general only extreme alternations with short periods at the higher temperature are effective in the dark. In the light any temperature alternation within the range 1–35°C is effective to at least some extent, providing the temperature difference is 5°C or more and providing the alternation includes one temperature which is above approximately 15°C and one which is below approximately 25°C. The optimum temperature difference is about 15°C. In the light, 4 to 10 cycles saturate the response, but in the dark, where the effect is much less, the response may not be saturated even by 16 cycles. KNO₃ at 10⁻³ M has little effect on the response to alternating temperatures either in the light or the dark. The response to alternating temperature regimes does not appear to vary in quality, i.e., in terms of which particular treatments are best, but it varies in magnitude with site and year of seed collection; and it increases slowly during dry storage, even when stored at a temperature as low as 1.5°C.     

Ontogenetic variation in response to temperature change in the control of seed dormancy of Rumex obtusifolius L. and Rumex crispus L.

Abstract After the onset of imbibition, the dormant seeds of Rumex obtusifolius and R. crispus are stimulated to germinate by a change from an initial low temperature to a warmer temperature for a relatively brief period: the warmer that temperature the shorter is the optimum period spent at it, and this optimum value is unaffected by the initial temperature. The optimum period is more critical in R. crispus than in R. obtusifolius (about 1 h and 2.5 to 4 h, respectively, for a warmer temperature of 35°C in the dark); in the light the length of the period at the warmer temperature is less critical in both species. The sensitivity of the seeds to the change to the warmer temperature increases with time from the start of imbibition at a rate which is positively related to the initial temperature. In R. obtusifolius maximum sensitivity was typically reached after 3 to 5 d when the initial temperature was 20°C and then remained constant, or declined only slightly, over the period investigated (10 d). At the same initial temperature, however, R. crispus showed a cyclical pattern of sensitivity with peaks occurring at 3–4 d intervals from the start of imbibition.     

The influence of the rate of temperature change on the activation of dormant seeds of Rumex obtusifolius L.

1. One temperature shift from 20 to 30°C in darkness induces 30–40% germination in Rumex obtusifolius seeds. The same germination percentages are found with heat treatment varying between 1 and 6h duration, indicating that the total heat sum of the temperature shift is not important.
2. Germination is greatly enhanced by three consecutive heat shifts of 1h at 30°C separated by 1h periods at 20°C.
3. The seeds are activated to a small extent after a slow warming (+2°Ch^–1) from 20 to 30°C, followed by incubation for 1h at 30°C. Germination is much higher after rapid heating (+10°Ch^–1) to 30°C, followed by 1h incubation at this temperature. Repeated fast heating treatments on four consecutive days enhances germination. Moderately rapid heatings (+3·3°Ch^–1) give intermediate results.
4. The rate of cooling does not influence the germination percentage. Cooling alone cannot induce germination.
5. Heating alone from 15 to 25°C without cooling also activates germination. In this temperature range the seeds are more activated by rapid warming than by slow warming.
6. The ecological relevance of the response to different warming rate is discussed. The insensitivity of seeds to a slow warming might keep deeply buried seeds in a dormant stage.     

Interaction of light and temperature on the germination of Rumex obtusifolius L.

Seeds (nutlets) of Rumex obtusifolius L. fail to germinate in darkness at 25° C, but are stimulated by short exposure to red light (R) the effectiveness of which can be negated by a subsequent short exposure to far red light (F) indicating phytochrome control. Short periods of elevated temperature treatment (e.g. 5 min at 35° C) can induce complete germination in darkness. Although short F cannot revert the effect of 35° C treatment, cycling the phytochrome pool by exposure to short R before short F results in reversion of at least 50% of the population. Prolonged or intermittent F can also revert the germination induced by 35° C treatment. The effect of elevated temperature treatment is interpreted on the basis of two possible models; (i) that it increases the sensitivity of the seeds to a low level of pre-existing active form of phytochrome (Pfr) (ii) that it induces the appearance of Pfr in the dark. In both cases it is envisaged that elevated temperature treatment and Pfr control germination at a common point in the series of reactions that lead to germination.Abbreviations D Dark - F far red light - P phytochrome - Pr red absorbing form of P - Pfr far red absorbing form of P - R red light     

Nitrate content and nitrate reductase activity in Rumex obtusifolius L.

Summary The aim of this work was to investigate the effect of nitrogen starvation and subsequent fentilization with nitrate or ammonium on nitrate content and nitrate reductase activity of Rumex obtusifolius L. under natural conditions.When plants were transplanted to nitrate-poor media, endogenous nitrate was reduced within a few days. In parallel, nitrage reductase activities dropped to about 25% of the initial values. As a consequence of nitrate fertilization (1; 10 or 100 mmol KNO₃/l substrate), endogenous nitrate content of the plant abruptly increased within one day. In extreme cases, nitrate concentrations of up to 10% of plant dry weight could be observed without being lethal. High external nitrate concentrations caused an inhibition of nitrate reductase within the leaves, while low external concentrations provoked an increase in the enzyme activity of about 450% within one day. Ammonium fertilization (5 mmol (NH₄)₂SO₄/l substrate) also caused an increase in nitrate reductase activity and nitrate content within leaf blades. This observation indicates a rapid nitrification of ammonium in the substrate. When plants were fertilized with ammonium plus nitrate (2.5 mmol (NH₄)₂SO₄+ 5 mmol KNO₃/l substrate), an extremely high and long term increase in nitrate reduction could be observed. Due to an intensive enzymatic nitrate turnover, the nitrate content of leaf blades then remained relatively low. Our observations do not point to an inhibition of nitrate reductase activity in leaves of Rumex obtusifolius by ammonium. Despite temporarily high endogenous nitrate concentrations, Rumex obtusifolius may not be termed as a nitrate storage plant, since the accumulation of nitrate is a short term process only.     

Nitrate content and nitrate reductase activity in Rumex obtusifolius L.

Summary With Rumex obtusifolius L., the influence of some environmental conditions on nitrate uptake and reduction were investigated. Nitrate concentrations of plant material were determined by HPLC, the activity of nitrate reductase by an in vivo test. As optimal incubation medium, a buffer containing 0.04 M KNO₃; 0.25 M KH²PO₄; 1.5% propanol (v/v); pH 8.0 was found. Vacuum infiltration caused an increase of enzyme activity of up to 40%.High nitrate concentrations were found in roots and leaf petioles. Nitrate reductase activity of these organs, however, was low. On the other hand, the highest nitrate reductase activity was observed in leaf laminae, which contained lowest nitrate concentrations.In leaves, nitrate content and nitrate reductase activity exhibited inverse diurnal fluctuations. During darkness, decreasing activities of the enzyme were followed by increasing nitrate concentrations, while during light the contrary was true. In petioles diurnal fluctuations in nitrate content were observed, too. No significant correlations with illumination, however, could be found.Our results prove that Rumex obtusifolius is characterized by an intensive nitrate turnover. Theoretically, internal nitrate content of the plant would be exhausted within a few hours, if a supply via the roots would be excluded.     

Interaction of light and temperature on seed germination of Rumex obtusifolius L.

Germination of Rumex obtusifolius L. seeds (nutlets) is low in darkness at 25° C. Germination is stimulated by exposure to 10 min red light (R) and also by a 10-min elevation of temperature to 35° C. A 10-min exposure to far-red light (FR) can reverse the effect of both R (indicating phytochrome control) and 35° C treatment. Fluence-response curves for this reversal of the effect of R and 35° C treatments are quantitatively identical. Treatment for 10 min with light of wavelenght 680, 700, 710 and 730 nm, after R and 35° C treatment, demonstrates that germination induced by 35° C treatment results from increased sensitivity to a pre-existing, active, far-red-absorbing form of phytochrome (Pfr) in the seeds.Abbreviations FR far-red light - P phytochrome - Pr red-absorbing form of P - Pfr far-red-absorbing form of P - R red light     

Performance and Mortality of Rumex obtusifolius and R. crispus in Managed Grasslands are Affected by Nutrient Availability

The reasons for the greater incidence of Rumex obtusifolius in grasslands compared with R. crispus have never been investigated. In a small-plot field experiment in which seedlings of R. obtusifolius and R. crispus were transplanted into a sward dominated by Dactylis glomerata, the growth and survival of seedlings were monitored over three years under control and P, N, NP and NPK fertilizer treatments. The highest plants of R. obtusifolius and R. crispus were generally recorded in the N, NP and NPK treatments and the lowest in the control and P treatment. The maximum recorded heights of R. obtusifolius were 100?cm and of R. crispus 80?cm. The number of leaves per plant, weight of individual plants, cover and fertility were generally higher for R. obtusifolius than for R. crispus. 30% to 80% of transplanted R. obtusifolius plants flowered in the first (seeding) year and this contrasted with no flowering of transplanted R. crispus plants. After cutting, substantially better regrowth of R. obtusifolius was recorded compared to that of R. crispus and surviving R. crispus plants tended to show a gradual reduction in leaf number. The mortality of R. obtusifolius over three years ranged from 0% in the NPK treatment to 13% in the control, with an average of 8% over all treatments. The mortality of R. crispus ranged from 19% in the control to 94% in the NPK treatment with an average of 64% over all treatments. We concluded that R. obtusifolius is better adapted to growing in highly productive temperate grasslands than R. crispus because of its perennial character, larger plant size and higher fertility.     

The effect of light on endogenous cytokinin levels in seeds of Rumex obtusifolius

Summary Exposure of Rumex obtusifolius seed to red light resulted in a rapid increase in extractable cytokinins. This increase can be reversed if the red light treatment is immediately followed by far-red irradiation. In germinating seed hardly any cytokinin activity could be detected in the butanol and aqueous extracts. An increase in the amount of activity present in the petroleum ether extracts of these seeds was, however, observed.The present findings suggest that cytokinin activity in light-sensitive seeds could be under the control of phytochrome.     

Characterization of glucosylceramides in the Polygonaceae, Rumex obtusifolius L. injurious weed

Rumex obtusifolius L., a member of Polygonaceae, is one of the world's worst weeds. We characterized the glucosylceramide molecular species in leaves of R. obtusifolius by liquid chromatography/tandem mass spectrometry. 4,8-Sphingadienines were principally paired with 2-hydroxy palmitic acids. In contrast, 4-hydroxy-8-sphingenines were chiefly attached to 2-hydroxy fatty acids with 22 to 26 carbon-chain length. A unique characteristic of the 2-hydroxy fatty acid composition of R. obtusifolius was the high content of n-9 monoenoic 2-hydroxy fatty acids with 22 and 24 carbon-chain length. The levels of the Z and E stereoisomers of the 8-unsaturated long-chain bases were reliably distinguished from those in other plant families in ten species of Polygonaceae.     

Interactions of seed water content with phytochromeinitiated germination of Rumex crispus (L.) seeds

Light-initiated germination levels of Rumex crispus L. seedswere reduced equally by imbibition in mannitol or polyethyleneglycol 6000 (PEG 6000) solutions of the same w, indicating thatthe effects of each were through w. Reduction of the water contentof the seeds with these osmotica decreased the effectivenessof the far-red absorbing form of phytochrome (P_tr) in causinggermination. However, reduced water content had no effect onthe slopes or saturation points of fluence/response curves whichindicates that has no effect on the number of P_tr receptor sites.The time during which a portion of the seeds were still photoreversibleby far-red light was increased by imbibition in PEG 6000, indicatinga direct effect of w on a reaction involving phytochrome. Noqualitative effect of PEG 6000 on the Onset of secondary dormancywas seen; however, its effect on the relative rate of appearanceof secondary dormancy was equivocal. ¹ Mississippi Agricultural and Forestry Experiment Station cooperating. (Received February 23, 1978; )     

Effects of added nitrogen on leaf nitrogen concentration and population dynamics of insects associated with Rumex obtusifolius L.

A series of experiments was designed to study the effects of changes in leaf nitrogen (N) concentration on population dynamics of a range of insects associated with Rumex obtusifolius. During the study period, the N concentration in leaves fluctuated between 2 and 6% dry weight. The insects at both study sites showed a pattern of shift in time for peak population density (time‐shift). This suggests that the insect community dynamics might be closely related to the allocation and distribution of resources by the host‐plant. The elevated N concentration in the host‐plant did not change the temporal order of the peak population densities of the insects, although it may determine the time for the optimal exploitation of resources by each insect species.     

Phytochrome Transformation and Action in Seeds of Rumex crispus L. during Secondary Dormancy

Promotion of germination by red light fails after prolonged dark imbibition of Rumex crispus L. seeds, indicative of a secondary dormancy. The degree and rate of inception of the dormancy increases with increasing temperature. Following establishment of the dormancy, germination response to red light can be restored by either prolonged cold treatment or brief high temperature shifts. Loss of phytochrome was not a factor in the initial establishment of the dormancy. When the seeds are in secondary dormancy, the chromophore of phytochrome can be transformed to the far red-absorbing form, but the far red-absorbing form cannot induce germination. The responses to changes in temperature suggested dependence of germination on order disorder transitions in components of the seeds.     

Photosensitivity of Rumex obtusifolius seeds for stimulation of germination: Influence of light and temperature

A population of Rumex obtusifolius L. seeds imbibed for 24 h at 25°C exhibits a sigmoid logarithmic fluence-response relationship for stimulation of germination by red light (R), 11.0 μmol m⁻² being necessary for 50% of the response. After 24 h imbibition at 35°C the fluence-response relationship for stimulation of germination by R is biphasic. For 50% response the very sensitive phase (very low fluence-response) requires 4.7 − 10⁻²μmol m⁻² whereas the less sensitive phase (low fluence-response) requires 4.0 μmol m². A few seconds of far-red light (FR) satisfies the germination requirement of the sensitive seeds after 24 h at 35°C. However, a longer period of FR (2 h) results in low germination. The fluence-response relationship for induction of these seeds by R is sigmoid, 4.8 μmol m⁻² being necessary for 50% response, demonstrating that 2 h FR desensitizes the sensitive proportion of the seed population induced by 24 h at 35°C. A proportion of the seed population can be further sensitized by 60 min at 35°C following this desensitization.     

Prevention of Action of Far-Red-Absorbing Phytochrome in Rumex crispus L. Seeds by Ethanol

Phytochrome-enhanced germination of curled dock (Rumex crispus L.) seeds is further stimulated by pretreatments in solutions of 0.5 to 2 molar methanol and 0.03 to ≥ 0.3 molar 2-propanol during a 2-day 20°C imbibition. Similar pretreatments in 0.1 molar ethanol, acetaldehyde, and n-propanol inhibit phytochrome-enhanced germination. If exposure to ethanol is delayed until 16 hours after a red irradiation, seeds escape the ethanol inhibition indicating a mechanism other than toxicity. The rate of escape from ethanol inhibition roughly parallels the escape from phytochrome control in seeds held in water only, indicating possible ethanol effects on phytochrome. It was found that ethanol pretreatment prevents the far-red absorbing form of phytochrome (Pfr) from acting but does not accelerate dark decay or prevent transformation. Ethanol inhibition may be prevented if ethanol pretreatment is at 10°C instead of 20°C, or may be overcome by transferring ethanol-pretreated seeds to 10°C in water. Similarly, ethanol inhibition can be overcome by a 2-hour 40°C temperature shift concluding the pretreatment. It is proposed that the ethanol causes perturbations at a membrane which prevent Pfr from acting.     

CO(2) Inhibits Respiration in Leaves of Rumex crispus L

Curly dock (Rumex crispus L.) was grown from seed in a glasshouse at an ambient CO₂ partial pressure of about 35 pascals. Apparent respiration rate (CO₂ efflux in the dark) of expanded leaves was then measured at ambient CO₂ partial pressure of 5 to 95 pascals. Calculated intercellular CO₂ partial pressure was proportional to ambient CO₂ partial pressure in these short-term experiments. The CO₂ level strongly affected apparent respiration rate: a doubling of the partial pressure of CO₂ typically inhibited respiration by 25 to 30%, whereas a decrease in CO₂ elicited a corresponding increase in respiration. These responses were readily reversible. A flexible, sensitive regulatory interaction between CO₂ (a byproduct of respiration) and some component(s) of heterotrophic metabolism is indicated.     

Phytochrome Control of Germination of Rumex crispus L. Seeds Induced by Temperature Shifts

High germination of curly dock (Rumex crispus L.) seeds is evident after suitable imbibition and temperature shift treatment, but germination at constant temperatures fails without an input of far red-absorbing form of phytochrome. Preliminary imbibitions at high temperatures (30 C) sharply reduce germination induced by temperature shifts. High germination may be restored by low energies of red radiation, or by brief far red adequate for the photosteady state. Prolonged far red during imbibition also nullifies temperature shift-induced germination. After prolonged far red, high germination may be restored by red radiation of an energy dependent upon the duration of the far red treatment. The evidence supports the conclusion that dark germination induced by temperature shifts arises from the interaction of pre-existent far red-absorbing form of phytochrome in the mature seeds with the temperature shift.     

Impacts of fire cues on germination of Brassica napus L. seeds with high and low secondary dormancy

• Agricultural burning is used in farm management operations; however, information about the impact of fire cues on the release and/or induction of secondary dormancy in crop seeds is scarce.
• Seeds from two oilseed rape cultivars were induced for high (HD) or low (LD) secondary dormancy using polyethyleneglycol (PEG) pre‐treatment, and their germination after exposure to various fire cues was compared to control PEG pre‐treated and non‐dormant seeds.
• Non‐dormant seed germination was unaffected by various fire cues. Low doses of aerosol smoke released secondary dormancy in HD seeds, while higher doses increased dormancy of LD seeds. Dilute smoke water also released HD seed secondary dormancy, but concentrated smke water enhanced dormancy in both LD and HD seeds. The concentrated aqueous extracts from charred oilseed rape straw only promoted germination of HD seeds, while dilution inhibited LD seed germination. Heat shock (80 °C, 5 min) released secondary dormancy in HD seeds; however, higher temperatures and/or increased exposure time was associated with seed death. GC‐MS analyses of smoke water revealed two butenolides and an array of monoaromatic hydroxybenzene compounds with potential germination inhibitor or promoter activity.
• The extent of secondary dormancy induction in seeds affects their subsequent responses to fire cues. Both aerosol smoke and smoke water have both germination promoter and inhibitor activity. Lacking any butenolides, aqueous extracts of charred straw contain a potential germination stimulating steroid, i.e. ergosterol. The significance of fire‐derived cues on behaviour of oilseed rape seeds in the soil seed bank is discussed.