Further Study on the Involvement of Cyanide-Resistant Respiration in the Germination of Cocklebur Seeds

The effect of propyl-gallate (PG) and benzohydroxamic acid (BHAM),inhibitors of cyanide-resistant, alternative respiration path(AP), on germination were examined using after-ripened upperand lower cocklebur (Xanthium pennsylvanicum Wallr.) seeds pre-soakedat 23?C for various periods. Germination was strongly suppressedby PG or BHAM at concentration above 2 mM. However, germinationwas enhanced by low concentrations of PG or BHAM (0.25 or 0.5mM) which reduced some portions of AP operation. Similarly,the high temperature-induced germination of pre-soaked upperseeds was promoted by the same low concentration range of PGor BHAM, in which PG and BHAM were effective only when appliedat the start of high temperature incubation. The inhibitionof germination by C₂H₄ at high temperature occurred only whenseeds were exposed to C₂H₄ during the earlier period of hightemperature incubation, and delayed application tended to promotetheir germination, although most of germinated seeds did notexhibit the normal germination behaviour of predominant radicleprotrusion. If the upper seeds had been subjected to a short-timepre-soaking, the inhibition of high temperature-induced germinationby C₂H₄ was prevented by the low concentrations of PG or BHAM,although the germination restored was mostly an abnormal, predominantlycotyldonary growth, suggesting that the germination inhibitionby C₂H₄ may be involved in some step of axial growth or in thegrowth of some specific axial zone. The lower concentrationsof PG or BHAM were promotive to the axial growth even at 33?C.Based on these results, the involvement of AP in cocklebur seedgerminaton is discussed in relation to the differential growthof axial and cotyledonary tissues. (Received May 2, 1986; Accepted October 27, 1986)     

A bimodal germination response to temperature in cocklebur seeds. II. ATP and adenylate pool

Abstract. The mechanism involved in a bimodal germination-temperature response in pre-soaked cocklebur (Xanthium pennsylvanicum Wallr.) seeds was studied with special reference to adenylate metabolism. Exposure to either low (optimal at 8°C) or high (optimal at 34°C) temperature which was effective in inducing the germination of the seeds brought about the accumulation of ATP in them. The ATP level remained unchanged at temperatures around 23°C. Pretreatment with KCN, stimulating germination even at 23°C, subsequently increased the ATP content, total adenylate pool and energy charge (EC) in the axial tissue prior to germination above those of the untreated controls. The lower the treatment temperature, the greater the inhibitory effect of KCN on ATP formation. An increase in germination following an increasing duration of pre-soaking at 8°C was comparable to increasing both the ATP content and total adenylate pool of axes, but not the EC value. Similarly, changes in germination following an increased exposure duration at 8°C correlated with changes in ATP content rather than EC value in the axes. Unlike the case of chilling, an increase in ATP level in response to 34°C was greater in the early period of water imbibition, during which times its germination-stimulating effect appeared more striking than in the later period, and it occurred without a concomitant rise in EC value because of the increased supply of AMP. Such a supply of AMP was reduced in the presence of benzohydroxamic acid or propyl gallale, inhibitors of an alternative respiratory pathway. It was thus concluded that both low temperature, coupled with warm temperature, and high temperature, by itself, can induce seed germination by increasing the ATP level as well as the total adenylate pool, but not the EC value, in the axial tissue. Further, that increases in both the ATP level and the adenylate pool especially are required for seed germination to proceed, probably depending on the activities of the cytochrome and alternative respiration pathways, respectively.     

Light Actions in the Germination of Cocklebur Seeds II. Possible Origin of the Diversity of Light Responses in Seed Germination

In negatively photoblastic, lower seeds of cocklebur (Xanthiumpennsylvanicum Wallr.), the respective germination-inhibitingeffects of red (R) and far-red (FR) lights were found in theproximal and near-tip zones of the axial tissues. In contrast,the germination-stimulating effect of R in positively photoblastic,upper cocklebur seeds was manifested in the near-tip zone ofthe axes, the R effect being reversed when FR was given to thezone. The R-sensitive zone in the upper seeds, however, shiftedtowards the more proximal zone as the period of pre-soakingat low temperatures increased. This shift was accompanied bythe ability to germinate in the dark in the upper seeds. In the lower seeds, R inhibited axial growth in the near-tipzone, whereas FR inhibited it in the proximal zone. In contrast,axial growth in the near-tip zone of the upper seeds was promotedby R. In both seeds, light had little effect on the growth ofthe radicle tip. Pre-soaking at low temperatures induced dark-germinationby hastening the axial growth of the upper seeds, thus allowingthe upper seed to resemble the lower one. We therefore proposea hypothesis that explains the diversification of photoresponsesin seed germination. (Received August 7, 1984; Accepted December 24, 1984)     

Light Actions in the Germination of Cocklebur Seeds: III. EFFECTS OF PRE-TREATMENT TEMPERATURE ON GERMINATION RESPONSES TO FAR-RED LIGHT AND ON DARK GERMINATION IN THE RED LIGHT-REQUIRING UPPER SEEDS

Esashi, Y., Oota, H., Saitoh, H. and Kodama, H. 1985. Lightactions in the germination of cocklebur seeds. III. Effectsof pre-treatment temperature on germination responses to far-redlight and on dark germination in the red light-requiring upperseeds.—J. exp. Bot. 36: 1465-1477. Red light (R) responsiveness in R-requiring upper cocklebur(Xanthium pennsylvanicum Wallr.) seeds changed in differentpatterns during a soaking period at different temperatures.At temperatures above 23°C, the responsiveness increasedand then decreased. At lower temperatures (3–18°C),however, it continued to increase throughout an experimentalperiod. The lower temperatures caused germination in the subsequentdark at 33°C, regained the R responsiveness and acquiredthe dark germinability when subsequently exposed to 8°C,to an extent proportional to the duration of the chilling. Far-red (FR) was inhibitory to germination in an earlier soakingperiod at lower temperatures, but its effect gradually decresed,and finally turned promotive. The negative FR response was repeatedlycontrolled by the following R irradiation. However, the positiveFR response was enhanced by an immediate R irradiation, andFR/R reversibility occurred after the second FR. In contrastto the R responsiveness and dark germinability, the positivegermination response to FR was not induced by soaking at 3°C,in which the growth of the axial tissue as a photoreceptivesite did not occur at all. Similarly, it was not manifestedwhen the seeds soaked at 33°C were subsequently subjectedto 8°C. Key words: Cocklebur seeds, dark germination, far-red light, low temperature, red light, seed germination, Xanthium pennsylvanicum     

β-Glucosidase activities and HCN liberation in unimbibed and imbibed seeds, and the induction of cocklebur seed germination by cyanogenic glycosides

In many seed species, the major source of HCN evolved during water imbibition is cyanogenic glycosides. The present investigation was performed to elucidate the role of endogenous cyanogenic glycosides in the control of seed germination and to examine the involvment of β-glucosidase in this process. All seed species used here contained some activities of β-glucosidase already in the dry state before imbibition. in the decreasing order of Malus pumila, Daucus carota, Hordeum vulgare, Chenopodium album and so on. β-Gluosidase activity in upper and lower seeds of cocklebur (Xanthium pennsylvanicum Wallr.) decreased with imbibition, and in lower seeds the activity disappeared when they germinated. On the contrary, in caryopses of rice (Oryza sativa L. cv. Sasanishiki) β-glucosidase increased during imbibition, and this increase continued even after germination. β-Glucosidase in cocklebur seeds was more active in the axial than in the cotyledonary tissue. Amygdalin, prunasin and linamarin could all serve as substrattes for the β-glucosidase(s) from both cocklebur and rice. Amygdalin, prunasin and linamarin as well as KCN, were effective in stimulating the germination of upper cocklebur seeds. The seeds evolved much more free HCN gas when they were exposed to the cyanogenic glycosides than when the glycosides were absent. Moreover, the application of the cyanogenic glycosides or of KCN caused accumulation of bound HCN in the seeds. Carbon monoxide, which stimulated cocklebur seed germination only slightly, did not cause accumulation of bound HCN. We suggest that a balance between the cytochrome and the alternative respiration pathways, which is adequate for germination (Esashi et al. 1987. Plant Cell Physiol. 28: 141–150), may be brought about by the action of endogenous HCN; a large portion of which is liberated from cyanogenic glycosides via the action of β-glucosidase. In addition to the partial suppression of the cytochrome path and unlike carbon monoxide, the HCN thus produced may act to supply cyanide group(s) to unknown compounds necessary for germination.     

Germination Responses of Taraxacum platycarpum Seeds to Temperature

Abstract Germination responses of Taraxacum platycarpum seeds to temperature were examined under laboratory conditions to investigate the emergence-season choice mechanism of the seeds. Almost all the newly collected seeds were non-dormant. Under constant temperature conditions, maximum percentage germination (approximately 90%) was attained at temperatures 6–16°C, where simple linear relationships were observed between the temperature and the rates of germination, i.e. the reciprocals of the time taken to germinate by seed subpopulations with 10–80% germination. Thermal time required for germination of the subpopulations ranged from 600 Kh (degree Kelvin × hours) to 1500 Kh with a relatively constant base temperature of about 2.5°C. Lower limit temperature for germination was slightly below 6°C. Higher limit temperature for germination has the normal distribution with the mean ±SD of 19±2.5°C. Pre-exposure of imbibed seeds to temperatures higher than the higher limit temperature for germination, 25 and 30°C, had no effect on the germinability and the rate of germination at a circa-optimum temperature. Moist chilling treatment at 4°C caused an increase in the variation of germination rate within the seed population, but no evidence for dormancy-inducing or breaking effects was obtained.     

Germination responses of Reaumuria vermiculata to salinity and temperature

Reaumuria vermiculata is a xerohalophytic perennial dwarf shrub distributed in many gypseous and saline areas in southern Tunisia. A laboratory experiment was carried out to assess the effects of temperature and salinity on germination. The germination responses of the seeds in darkness were determined under a range of salinity and temperature regimes. Temperatures between 10°C and 30°C seem to be favourable for the germination of this species. Germination was inhibited by either an increase or a decrease in temperature from the optimal temperature (15°C). Highest germination percentages were obtained under non-saline conditions and increases in salinity inhibited seed germination. Salt stress decreased both the germination percentage and the germination speed (increase of the mean time to germinate). Seed germination decreased with an increase in NaCl concentrations at all temperatures. An interaction between salinity and temperature yielded no germination at salinity of 300 mM, whereas some seeds germinated under the optimal temperature.     

Light Actions in the Germination of Cocklebur Seeds: IV. DISAPPEARANCE OF RED LIGHT-REQUIREMENT FOR THE GERMINATION OF UPPER SEEDS SUBJECT TO ANOXIA, CHILLING, CYANIDE OR AZIDEPRETREATMENT

Esashi, Y., Fuwa, Nn Kojima, K. and Hase, S. 1986. Light actionsin the germination of cocklebur seeds. IV. Disappearance ofred light-requirement for the germination of upper seeds subjectto anoxia, chilling, cyanide or azide pretreatmenL—J.exp. Bot. 37: 1652–1662. The effects on the germination of positively photoblastic uppercocklebur (X anthium pennsylvanicum Wallr.) seeds by pretreatingwith anoxia, chilling, cyanide or azide, which stimulates theirdark germination, were examined in relation to light actions.Prior to experiments, seeds were pre-soaked at 23 °C inthe dark for 1 or 2 weeks to remove the pre-existing Pfr. Whenthe prctreatment conditions were suboptimal for germinationinduction, the stimulating effects of the pretreatments on germinationduring a subsequent dark period at 23 °C were manifest onlywhen seeds were irradiated with red light before or after thepretreatment Red light promotion was reversed by blue or far-redlight treatment. However, both prc-chilling for 6 d at 8 °Cand prctreatment with 1· 5 mol m ^{– 3} NaN³ for 2d could induce full germination without red light exposure.On the other hand, both pre-exposure to anoxia for 8 d and pretreatmentwith 30 mol m^–3 KCN could induce the dark germinationonly when germination occurred at 33 °C which is known toaugment the ratio of an alternative respiration flux to a cytochromeone. Moreover, the dark germination in response to these inductionswere strongly inhibited by the inhibitors of alternative respiration,propyl gallate and benzohydroxamic acid, applied during a subsequentdark period. It was thus suggested that Pfr has some relationto the operation of two respiration systems of cocklebur seeds,but it is not indispensable to germination of this positivelyphotoblastic seed. Key words: Anoxia, azide, blue light, chilling cyanide, dark germination, far-red light, red light, seed germination, X anthium pennsylvanicum     

Stimulation of ethylene production by hydroxamic acid, in particular, by benzohydroxamic acid

The effects on the ethylene production of known inhibitors ofa cyanide-insensitive, alternative respiration in plants wereinvestigated using cotyledonary segments of cocklebur (Xanthiumpennsylvanicum Wallr.) seeds. Benzohydroxamic acid (BHAM) at3 mM stimulated ethylene production 4- to 8-fold over the control,but respiration of the segments was hardly affected at thatconcentration. The stimulatory effects of 3-chlorobenzohydroxamicacid (CLAM) and salicylhydroxamic acid were far smaller thanthat of BHAM. BHAM at 3 mM also markedly stimulated the ethyleneformation in the epicotyl or hypocotyl sections of etiolatedpea (Pisum sativum L.) and mung bean (Vigna radiata [L.] Wilczek)seedlings. Moreover, 3 mM BHAM further promoted the increasedethylene formation which was caused by administration of 1-aminocyclopro-pane-1-carboxylicacid (ACC) to the cotyledonary segments. The promoting effectsby BHAM and CLAM were also found in the conversion of ACC intoethylene in pea stem homogenates. (Received July 26, 1980; )     

Stimulation of ethylene production by hydroxamic acid, in particular, by benzohydroxamic acid

The effects on the ethylene production of known inhibitors ofa cyanide-insensitive, alternative respiration in plants wereinvestigated using cotyledonary segments of cocklebur (Xanthiumpennsylvanicum Wallr.) seeds. Benzohydroxamic acid (BHAM) at3 mM stimulated ethylene production 4- to 8-fold over the control,but respiration of the segments was hardly affected at thatconcentration. The stimulatory effects of 3-chlorobenzohydroxamicacid (CLAM) and salicylhydroxamic acid were far smaller thanthat of BHAM. BHAM at 3 mM also markedly stimulated the ethyleneformation in the epicotyl or hypocotyl sections of etiolatedpea (Pisum sativum L.) and mung bean (Vigna radiata [L.] Wilczek)seedlings. Moreover, 3 mM BHAM further promoted the increasedethylene formation which was caused by administration of 1-aminocyclopro-pane-1-carboxylicacid (ACC) to the cotyledonary segments. The promoting effectsby BHAM and CLAM were also found in the conversion of ACC intoethylene in pea stem homogenates. (Received July 26, 1980; )     

Changes in dormancy of Sisymbrium officinaleseeds do not depend on changes in respiratory activity

Effects of dark incubation at different temperatures were studied on dormancy and respiratory activity of seeds of Sisymbrium officinale (L.) Scop. Because germination of this species absolutely depends on the simultaneous action of light and nitrate, changes in dormancy could be studied in darkness without the interference of early germination events. Upon the start of incubation rates of O₂ uptake and CO₂ release rose. This was followed by a gradual decrease until stable levels of O₂ uptake and CO₂ release were achieved. Seeds kept for prolonged periods at 24^°C, showed neither a change in germination capacity nor in rates of O₂ uptake and CO₂ release. Respiratory quotients were 0.55–0.7. The initial rise in O₂ uptake correlated with the rate of water uptake and with breaking of primary dormancy. However, the subsequent decline in O₂ uptake was not generally linked to induction of secondary dormancy. An increased O₂ uptake was not required during breaking of secondary dormancy. It is concluded that changes in dormancy are not generally related to changes in respiratory activity. However, germination strongly depends on respiration. The increase in O₂ uptake started well before radicle protrusion. A far red irradiation only reversed this increase when it was given before germination escaped from its red light antagonising action. The contribution of different respiratory pathways was followed during prolonged incubation at 24^°C in darkness. KCN at 1.5 mM was needed to inhibit the cytochrome pathway (CP) and benzohydroxamic acid (BHAM) at 30 mM to inhibit the alternative pathway (AP). These concentrations did not exert any side effects. Electron flow was predominantly via the CP, maximally 10% was via the AP. Flow through the CP declined during the first 6 days and residual respiration remained constant. Therefore, the contribution of residual respiration became relatively more important with prolonged incubation. KCN at concentrations that almost completely inhibited flow through the CP, did not dramatically reduce germination. BHAM already inhibited germination at concentrations that do not inhibit oxygen uptake.     

Thermoperiodism Mechanism in the Germination of Cocklebur Seeds

In thermoperiodic germination of secondarily dormant cocklebur(Xanthium pennsylvanicum Wallr.) seeds, the extent of electronflow through the CN-sensitive, cytochrome path and the CN-resistant,alternative path differred in the cool and warm phases. Thecool phase triggered active engagement of the alternative pathduring the subsequent warm phase, and this led to an increasein the ratio of the alternative path flux to the cytochromepath flux. The cool and warm phases had different functionsin adenylate metabolism. The former acted mainly in the productionand accumulation of ATP, whereas the latter supplied ADP and,especially, AMP. Thus an increasing number of sequential thermoperiodiccycles augmented the size of the adenylate pool and the energycharge, both of which may be necessary for germination to takeplace. (Received September 16, 1981; Accepted November 5, 1982)     

Rhythmic Behavior in Germination of Cocklebur Seeds

Non-dormant, lower seeds of cocklebur (Xanthium pensylvanicum Wallr.) germinated with unimodal flush after 20 and 36 h from the start of water imbibition at 33 and 23°C, respectively. At 28°C, however, germination occurred bimodally, the time of each peak coinciding with that at 23 and 33°C. This type of germination behavior was induced even at 33°C, when the seeds were contacted with some osmotica. Further, the application of different osmotica at 28°C caused a rhythmic multimodal germination with a period of about 16 h. It was suggested that an endogenous rhythmicity may be involved in the control of cocklebur seed germination.     

Interrelation between Ethylene and Carbon Dioxide in Relation to Respiration and Adenylate Content in the Pre-Germination Period of Cocklebur Seeds

Effects of C₂H₄ and CO₂ on respiration of pre-soaked upper cocklebur(Xanthium pennsylvanicum Wallr.) seeds during a pre-germinationperiod were examined in relation to effects of the two gaseson germination. At 33?C, cocklebur seed germination was greatlystimulated. This high temperature-stimulated germination wasseverely inhibited by C₂H₄, but not by CO₂, although both gasesstimulated germination at 23?C. C₂H₄ promoted seed respirationat 23?C, but its promotive effect decreases with increasingtemperature and disappeared at about 35?C, while CO₂ stimulatedrespiration regardless of temperature. CO₂ augmented the operationof the CN-sensitive, cytochrome path (CP) regardless of temperature,resulting in an increase in the ratio of the CP flux to a CN-resistant,alternative path (AP) flux. On the other hand, C₂H₄ augmentedthe operation of both paths, particularly of the AP, at 23?C,where it promoted germination. However, at 33?C where germinationis suppressed by C₂H₄, C₂H₄ preferentially stimulated respirationvia the AP, thus leading to an extremely high ratio of AP toCP. The inhibitory effect of C₂H₄ on germination at 33?C disappearedcompletely in enriched O₂, under which conditions CP is knownto be augmented. At 23?C, CO₂ and C₂H₄ acted independently incontrolling seed respiration, but they were antagonistic at33?C. The independent action appeared when the AP flux was verylow relative to the CP flux, while the antagonism appeared whenthe AP flux had risen. This differential action of the two gasesat different temperatures was also observed in the ATP level,adenylate pool size and energy charge of the axial tissues.These results suggest that the germination-controlling actionsof both CO₂ and C₂H₄ are fundamentally manifested through themodification of respiratory systems. However, the germination-inhibitingeffect of C₂H₄ at 33 ?C was not removed by inhibitors of AP,and there was little difference in the adenylate compounds betweenthe C₂H₄-treated and non-treated seeds at 33?C. Therefore, thephysiological action of C₂H₄ can not be explained only in termsof regulation of the respiratory system. (Received January 24, 1986; Accepted November 17, 1986)     

Temperature control of germination and its possible role in the survival of a non-dormant population of Avena fatua

Germination of freshly harvested seeds of a non-dormant (ND) line (Stonehouse 319) of wild oats ( Avena fatua L.) was inhibited by incubation of the seeds at relatively high temperatures of 25 and 30°C. The germination inhibition in these seeds appeared to be a case of thermo-inhibition which was the direct effect of hightemperature treatment (HIT), since it did not persist after transferring the seeds to an optimum germination temperature of 20°C. Even a prolonged HTT of 30°C for over 5 weeks did not prevent germination of about 80% of the seeds transferred to 20°C. However, in a significant proportion of the seeds, thermo-dormancy was induced by 10 days of HTT at 30°C if the seeds were then incubated at sub-optimal temperatures of 5 to 15°C. This thermo-dormancy would appear to be 'restrictive' in form, since its expression was restricted to very specific conditions. Relatively low inclubation temperaturs of 5 and 10°C markedly slowed germination whether HTT was applied or not. The results suggest that thermo-inhibition and thermo-dormancy, induced during seasonal temperature fluctuations, may provide a survival mechanism for seeds of such ND lines as Stonehouse 319.     

Cytochrome and alternative pathway respiration in white spruce (Picea glauca) roots. Effects of growth and measurement temperature

Interactions between growth temperature and measurement temperature were examined for their effects on white spruce [ Picea glauca (Moench) Voss] root respiration. Total dark respiration rates increased with measurement temperature and were unaffected by growth temperature. Partitioning of respiratory electron flow between the cytochrome and alternative pathways was also unaffected by growth temperature. The proportion of respiration mediated by the alternative pathway was constant at measurement temperatures between 4°C and 18°C, but was increased at higher temperatures. Changes in alternative pathway activity were paralleled by changes in capacity, and the alternative pathway was almost fully engaged at all temperatures. Roots grown at low temperature displayed higher carbohydrate levels than roots grown at higher temperatures, but respiration rate was unaffected. Spruce root respiration did not appear to acclimate to growth temperature, and the alternative pathway was not preferentially engaged at low temperature.     

Temperature sensitivity of in vitro germination and seedling development of Dactylorhiza majalis (Orchidaceae) with and without a mycorrhizal fungus

Abstract. Symbiotic germination and development in vitro of Dactylorhiza majalis seeds with a strain of Rhizoctonia is very temperature dependent. Above an optimum at 23–25 °C there is a marked decline in germination percentage. Seeds that did germinate at higher temperatures had only little or no development of mycorrhiza, and developed few or no rhizoids compared with seedlings raised at optimal or lower temperatures. Six-week-old seedlings grown for additional 4 weeks on a range of temperatures had an optimal length increase at 23–24.5 °C mean temperature. At superoptimal temperatures (26 °C), the seedlings contained smaller starch reserves than those at lower temperatures and increased about as much in length as seedlings grown at 13 °C but much less than those grown at optimum. Temperature also influenced the differentiation of the leafy shoot, seedlings growing to a larger size before shoot initiation in the temperature range of optimal growth. Because of the small span between optimal and too-high temperatures, a careful assessment of temperature optimum will be necessary in any orchid/fungus relationship before judging the success of symbiosis. At optimal temperature, symbiotic germination gave a germination percentage about twice that using a good asymbiotic method. The increase in seedling length was about 45% per week in symbiotic culture compared with less than 30% in the asymbiotic culture.     

Light and temperature action in germination of seeds of the empress tree (Paulownia tomentosa)

Seeds of the empress tree ( Paulownia tomentosa Steud.) were imbibed for two weeks in darkness at constant temperatures (18, 23 or 28°C), and then irradiated with red light for 5 min. Germination was poor if it took place at the same temperature as imbibition, but a high percentage was achieved if the seeds were exposed to higher or lower temperatures before they were irradiated. Maximum germination was obtained when the difference between pretreatment and imbibition was about 10°C. The effect increased with the duration of the pretreatment and was optimal at 24 h. The effect decreased as the time lapse between temperature pretreatment and red light irradiation increased, and it was lost after two days. If pretreatment was shorter than 24 h (12 h). a high percent of germination was obtained by alternating pretreatment and imbibition temperatures. The germination of seeds imbibed in 40% heavy water was also stimulated by temperature pretreatments. Light and temperature also exhibited an interactive effect in the germination of seeds that were imbibed in darkness for only 3 days. For each of the germination phases there was a temperature at which the time needed for 50% germination was the shortest, namely 35°C during imbibition, 37.5°C in the period of P_fr activity. and 32.5°C during radicle protrusion. The data obtained are shortly discussed in relation to the domestication of empress tree in Southern Europe.     

Developmental strategies of Koenigia islandica, a high-arctic annual plant

Seed germination, growth and flowering of the arctic-alpine annual Koenigia islandica were studied in controlled environment. Intact (unabraded) seeds germinated poorely at temperatures up to 18°C, with an optimum at 24°C (89% in 10 d). Scarified seeds germinated rapidly, and reached 100% germination in 3 d at 21°C, but no >40% germination occurred at 9 and 12°C, The seeds had no light requirement for germination, nor did fluctuating temperatures improve germination
Dry matter production was optimal at 12°C in both short day (SD) and long day (LD) conditions, but was markedly higher in LD than in SD at identical fluences at all temperatures except 21°C where the plants showed symptoms of severe heat stress. The temperature compensation point for net productivity was estimated to 24°C, and negative carbon balance at higher temperatures might be an important physiological mechanism limiting the distribution of K. islandica in Scandinavia.
Flowering was extremely rapid and independent of daylength, even in a high-arctic population from 79°N, In full summer daylight anthesis was reached 24 d after germination and seeds ripened after 36 d at 15°C, Days to anthesis varied little across the temperature range from 6 to 21°C, giving a linear decrease in the heat-sum requirement for the attainment of flowering with decreasing temperature.
It is concluded that conservative seed germination strategy, tininess and rapid development, low temperature optima for growth and reproduction, and daylength indifference of flowering are important adaptations for success of an annual plant in high-arctic and high-alpine environments, Daylength neutrality has facilitated the wide-latitudinal distribution of K. islandica. including the penetration of the species to the southern hemisphere.     

Changes in sensitivity of Amaranthus retroflexus L seeds to ethylene during preincubation. II. Effects of alternating temperature and burial in soil

Abstract. The effects of diurnally alternating temperatures and of prolonged burial in the soil on germination response of redroot pigweed ( Amaranthus retroflexus L.) seeds to ethylene were investigated. Percentage germination in a 12 h/12 h, 23° C/35° C temperature regime roughly equalled that observed at constant 35° C, and greatly exceeded that observed at 30°C. Preincubation for 61 d in alternating temperatures, which were gradually increased to simulate soil warming in spring, caused little germination in the absence of ethylene, but considerably enhanced sensitivity to ethylene. Seeds kept in soil in the same temperature regime failed to show the response to ethylene, and the soil itself removed ethylene from the soil atmosphere.
After burial in a field plot either over winter or during the summer, seeds had a very low ethylene response threshold (0.01−0.05 cm³ m⁻³) and strong response to ethylene (70–95% germination at 51 cm³ m⁻³ compared to 1–20% without ethylene). Germinability of seeds buried overwinter declined between 10 May (85%) and 24 May (7%), and 90% of those recovered on or after 24 May had a visible rupture in the seed coat. Apparently, germination had begun during burial, but was arrested by unknown causes in an early phase and was followed by seed deterioration.
Although the role of ethylene in germination of buried seeds remains uncertain, the greatly enhanced sensitivity to ethylene observed in pigweed seeds after burial deserves further investigation.