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)     

Cyanide-sensitive and Cyanide-resistant Respiration in the Germination of Cocklebur Seeds

Interrelation between the CN-sensitive cytochrome path and the CN-resistant, benzohydroxamic acid (BHM)-sensitive, or n-propylgallate (nPG)-sensitive alternative path in seed respiration during germination was examined using the nondormant upper and lower seeds of Xanthium pensylvanicum Wallr. The operation of both paths was required not only for normal germination of the lower seed but also for KCN- or NaN₃-induced germination of both. From the sensitivity to BHM of the germination response, it became obvious that the alternative path exerts its physiological activity as soon as it develops during the early period of water imbibition. Pretreatments with KCN and NaN₃ for promoting germination, strikingly decreased only the engagement of the cytochrome path in the subsequent respiration without affecting that of the alternative path. Nevertheless, no germination occurred without the operation of the cytochrome path. This suggested that excess operation of the cytochrome path is detrimental to germination, being maximal following the BHM-sensitive phase.     

Possible Involvement of the Alternative Respiration System in the Ethylene-stimulated Germination of Cocklebur Seeds

Respiration of nondormant upper cocklebur (Xanthium pensylvanicum Wallr.) seeds was enhanced by exogenous C₂H₄, proportionally to the concentration of C₂H₄ and the duration of presoaking of the seeds. Benzohydroxamic acid (BHM) and salicylhydroxamic acid (SHM), inhibitors of alternative respiration, inhibited both the germination of nondormant lower cocklebur seeds and the respiration of the upper seeds presoaked for periods of 12 to 30 hours. Both the growth and respiration of axial and cotyledonary tissues were also inhibited by BHM. Moreover, BHM inhibited both the C₂H₄-induced germination of the upper seeds and their C₂H₄-stimulated respiration; the inhibition occurred only with concomitant addition of C₂H₄ and BHM. The respiration of seeds with a secondary dormancy induced by presoaking for prolonged periods was markedly stimulated by C₂H₄ but not suppressed by BHM. It was suggested that the alternative respiration system may be involved in the normal germination process of cocklebur seeds, secondary dormancy may result from its inactivation, and C₂H₄ may exert its germination-promoting action by stimulating the alternative respiration. The effects of BHM and SHM can suggest but not prove the involvement of the alternative respiration in seed germination.     

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)     

Inhibitory Effects of Methyl Jasmonate on the Germination and Ethylene Production in Cocklebur Seeds

Methyl jasmonate (JA-Me) inhibited the germination of cocklebur (Xanthium pennsylvanicum Wallr.) seeds. The inhibition of the germination of cocklebur seeds treated with JA-Me at concentrations less than 300 μm was nullified by ethylene applied exogenously, although the inhibitory effect of 1,000 μm JA-Me was not recovered completely even by high concentrations of ethylene (10,000 μL/liter). JA-Me inhibited ethylene production before seed germination. The level of 1-aminocyclopropane-1-carboxylic acid (ACC) in the cotyledonary tissues treated with JA-Me decreased but not the level of 1-(malonylamino)cyclopropane-1-carboxylic acid (MACC). JA-Me inhibited the conversion of ACC to ethylene in the tissues. These results suggested that JA-Me inhibits ethylene production by prevention of ACC oxidation in addition to ACC synthesis. We believe that the inhibition of ethylene production by JA-Me results in the retardation of the germination of cocklebur seeds. Received June 4, 1997; accepted October 23, 1997     

Necessity of a Balance between CN-Sensitive and CN-Resistant Respirations for Germination of Cocklebur Seeds

When applied singly, KCN or NaN3, as inhibitors of the cytochromerespiration path, and benzohydroxamic acid or n-propyl gallate,as inhibitors of the alternative respiration path, were lesseffective and ineffective, respectively, in inducing germinationof secondarily dormant, upper seeds of Xanthium pennsylvanicumWallr. When applied in combination, however, these chemicalswere very effective, producing much higher gemination. Thus,we concluded that an appropriate balance between the cytochromeand alternative path fluxes is required to induce the germinationof secondarily dormant cocklebur seeds. (Received August 30, 1980; Accepted December 2, 1980)     

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)     

Dormancy and Impotency of Cocklebur Seeds VIII. Lack of Germination Responsiveness in Primarily Dormant Seeds to Cyanide, Azide, Anoxia and Chilling

Germination responsiveness to KCN, NaN₃, chilling or anoxiaand respiration activity was compared between non-after-ripenedand after-ripened upper cocklebur (Xanthium pennsylvanicum Wallr.)seeds. The latter, coat-imposed dormant seeds, could germinatein response to the above chemicals and conditions, whereas theformer, primarily dormant seeds, could not respond. There waslittle difference in the respiratory properties of both typesof seeds. (Received June 22, 1981; Accepted October 30, 1981)     

Ethylene Production in Pea and Cocklebur Seeds of Differing Vigour

Relationships between seed vigour and ethylene (C₂H₄) productionwere studied using C₂H₄-responsive fatty cocklebur seeds (Xanthiumpennsyhanicum Wallr.) and C₂H₄-insensitive starchy pea seeds(Pisum sativum L. cv. Alaska), which had been harvested in differentyears and subjected to different storage conditions. In bothspecies, the seeds with the highest vigour evolved the largestamounts of C₂H₄ during a period of water imbibition. The reductionof C₂H₄ production in cocklebur seeds occurred concomitantlywith the reduction in the growth potentials of both axial andcotyledonary tissues. Similarly, the activity of ACC-C₂H₄ conversionincreased with soaking, and was greater in seeds of high vigourcompared with those of low vigour. However, the change in ACCcontent in pea seeds differed from that in cocklebur seeds.That is, pea seeds with high vigour accumulated less ACC duringan imbibition period than those with low vigour. From theseresults it was suggested that the inferior C₂H₄ production bylow vigour pea seeds is mainly attributable to low ACC-C₂H₄conversion, whereas that by low vigour cocklebur seeds is dueto the shortage of ACC supply in addition to the reduced ACC-C₂H₄conversion. However, germination of deteriorated cocklebur seedswas not restored by exposure to ACC or C₂H₄, suggesting thatthe loss of seed vigour reduces the responsiveness of seedsto C₂H₄ as well as C₂H₄ production. Key words: Pea, cocklebur, seed vigour, ethylene production, 1-aminocyclopropane-1-carboxylic acid     

Identification and Content of 1-Malonylaminocyclopropanecarboxylic Acid in Germinating Cocklebur Seeds

A major conjugate of 1-aminocyclopropanecarboxylic acid in germinatingcocklebur (Xanthium pennsylvanicum Wallr.) seeds was isolatedand identified as 1-malonylaminocyclopropanecarboxylic acid(MACC). The change in MACC content during the germination periodof this seed also was examined. (Received November 4, 1983; Accepted March 15, 1984)     

棕榈种子萌发特性及其贮藏行为的研究

对3个不同种源地的棕榈(Trachycarpus fortunei)种子进行了不同时间的脱水处理和两种不同温度(4℃和-20℃)下贮藏(120d),对其萌发特性和贮藏行为进行了比较分析,结果表明棕榈种子脱水耐性低和对低温较敏感,且含水量较低的种子也受低温伤害。经不同含水量与温度的组合试验后发现,已实验的棕榈种子极可能是一种中间性种子,且其特性受其自然生境的深刻作用,尤其是海拔高度的影响。     

Light Actions in the Germination of Cocklebur Seeds: I. DIFFERENCES IN THE LIGHT RESPONSES OF THE UPPER AND LOWER SEEDS

Germination responses to light were studied in the upper andlower seeds of cocklebur (Xanthium pennsylvanicum Wallr.). Thelower seed was dark-germinating and negatively photoblastic;the upper one had a red-light (R) requirement and was positivelyphotoblastic. Germination of the lower seeds was inhibited bya prolonged single irradiation with R, blue (B) or far-red (FR)light applied during imbibition. The maximal inhibitory effectof a single irradiation occurred 9 h and 13 h after the startof soaking at 33 °C and 23 °C, respectively. However,the inhibitory effect of R differed from that of B and FR, byonly delaying germination. A single exposure to B or FR lightcould be replaced by intermittent B or FR irradiation, and theireffects were repeatedly reversible by the following R irradiation.If the upper seeds were not exposed to R during imbibition,they failed to germinate even at 33 °C which was optimalfor germination, and the promotive effect of R increased withdelay of its application time. The photoperceptive locus incocklebur seeds was the axial tissue for all B, R and FR. Lightreceived by the cotyledonary tissue had little effect. Germinationdimorphism in response to light is discussed with respect tothe phytochrome content and the ageing of axial tissues. Key words: Blue light, Dimorphism, Far red light, Germination, Red light, Xanthium seed     

Involvement of Cyanogens in the Promotion of Germination of Cocklebur Seeds in Response to Various Nitrogenous Compounds, Inhibitors of Respiratory and Ethylene

Nitrogenous inhibitors of respiration, namely, KCN, NaN₃ andNH₂OH, which promote the germination of cocklebur (Xanthiumpennsylvanücum Wallr.) seeds, enhanced the accumulationof cyanogenic compounds. Urea and cyanamide, which were ineffective,did not. Part of the exogenously applied KCN was converted tocyanogenic glycosides and lipids, but these compounds were onlyproduced at low levels in NaN₃-treated seeds. Exposure to NH₂OHcaused a slight accumulation of both types of cyanogenic compounds.Thiourea, effective in stimulating germination, did not increasethe cyanogenic compounds, which suggests that the mechanismof its effect on germination is different from that of othernitrogenous compounds. Thiocyanate had a slightly promotiveeffect and caused minor increases in the levels of cyanogeniccompounds. Ethylene increased the metabolic utilization of the accumulatedcyanogens in response to KCN or NaN₃. The response to ethyleneof seeds in secondary dormancy was restored only after pre-treatmentwith KCN or NaN₃. The response occurred once the level of cyanogenshad increased. By contrast, enhancement of the effects of KCNand NaN₃ on germination by propyl gallate or benzohydroxamate,inhibitors of CN-resistant respiration, was accompanied by theincreased accumulation of cyanogens within the seeds that hadbeen exposed to KCN or NaN₃ in combination with the other typesof respiratory inhibitors. Thus, it is suggested that endogenouscyanogens might be involved in the germination of cockleburseeds. (Received December 20, 1995; Accepted April 9, 1996)     

Failure of Chlorophyll Formation in the Cotyledons from Secondarily Dormant Cocklebur Seeds

The cotyledons of secondarily dormant cocklebur (Xanthium pennsylvankumWallr.) seeds lacked not only growing potential but also chlorophyll-formingability. These properties developed as the seeds establishedsecondary dormancy during a soaking period. The inability toform chlorophyll in cotyledonary segments was not improved inthe presence of S-amino levulinic acid. But this ability wasgreatly restored by benzyladenine or ethylene treatment, whichis effective in increasing cotyledonary growth. Similarly, theapplication of ethylene together with enriched oxygen and carbondioxide, or subsequent to a KCN treatment, which were the mosteffective means for breaking secondary dormancy, completelyrestored both chlorophyll formation and growing ability of thecotyledons. The depression of the greening-ability in cotyledonsmay be involved in development of secondary dormancy of cockleburseeds in concert with the decline of their growth pontential. (Received September 24, 1982; Accepted December 27, 1982)     

Germination Preventing Mechanisms in Iris Seeds

The different germination behaviour of the seeds of two irises,Iris lorteti and I. atropurpurea was found to be due to thedifferent mechanical resistance of the integument, at the micropylarend, to radicle protrusion. A pressure of 135 atm was necessaryin l. lorteti seeds for radicle protrusion, while in I. atropurpurea77 atm was sufficient. In contrast Pancratium maritimum requireda pressure of only 10 atm. The outer integument of seeds ofI. lorteti was found to contain a compound which was toxic tothe germinated embryo but did not act as a germination inhibitor.Extracts of the endosperm also had a slight germination inhibitingeffect. An interaction between this weak inhibitor and the effectof the testa could not be ruled out completely. A test assayfor germination using excised embryos was developed. A methodfor germination of Iris seeds, by cutting off the outer integumentat the micropylar end, was developed and is being exploitedcommercially. Iris lorteti, Iris atropurpurea, germination, germination inhibition, embryo culture, seed coat mechanical resistance     

Germination Inhibitors in some Umbellifer Seeds

Presence of inhibitors of germination in seeds of five umbellifercrops viz. fennel (Foeniculum vulgare Mill.), cumin (Cuminumcyminum L.), carum (Carum copticum Benth. and Hook.), carrot(Daucus carota L.), and coriander (Coriandrum sativum L.) wasdemonstrated. The effect of seed exudates on the pattern ofinhibition of germination of these seeds was also studied. Presence of exudates in general delayed germination althoughthe time of maximum inhibition was different with differentcrop seeds. The inhibitory effect was high at an early stagein the case of carum and carrot and persisted in the latterbut disappeared in the former. The inhibitory effect on cuminand fennel increased up to the seventh day and then began todecrease. Growth of root and hypocotyl of 16-day-old seedlingswas also inhibited by the exudates.     

Promotion of Cocklebur Seed Germination by Allyl, Sulfur and Cyanogenic Compounds

The effects of allyl, sulfur and cyanogenic compounds on thegermination of upper cocklebur (Xanthium pennsylvanicum Wallr.)seeds were examined. Mercaptoethanol and methylmercaptan aswell as KCN, substrates for rßcyanoalanine synthase(CAS), and H₂S and thiocyanate, the products of the CAS catalyzingreaction, were effective in promoting germination, suggestingthe involvement of CAS in germination. Most of allyl compounds, especially allylthiourea, as well asethylene which activated CAS [Hasegawa et al. (1994) Physiol.Plant. 91: 141], promoted the germination in an abnormal typewhich occurred by the predominant growth of cotyledons as didC₂H₄ [Katoh and Esashi (1975) Plant Cell Physiol. 16: 687].However, they failed to activate CAS unlike ethylene, and toliberate free ethylene during an incubation period. It was thuspossible that an C₂H₄-like double bond within allyl compoundscan act to promote seed germination. (Received June 10, 1996; Accepted August 21, 1996)     

葡萄属种子发育的物候、萌发行为及其对冷层积的反应

以3个起源和分布中心的不同葡萄种及其杂交种作为实验材料, 观察了开花、果实和种子成熟的物候, 研究了成熟种子的基本特性和最初萌发率以及冷层积对成熟种子萌发的影响。结果表明, 葡萄种子具有休眠特性, 生理成熟时都具有分化完全的胚, 在休眠解除过程中胚的形态不发生变化; 美洲种休眠最深, 欧亚种休眠最浅, 其它种类的葡萄种子的休眠程度介于美洲种和欧亚种之间; 不同种葡萄花期、果实和种子成熟过程的物候存在差异, 果实成熟与种子成熟不同步, 其间隔时间越短, 种子休眠程度越深。冷层积60天能有效地解除东亚种、欧亚种、欧山杂种和蘡欧杂种的种子休眠, 但对解除美洲种、美洲种种间杂种和欧美杂种的种子休眠效果较差。     

Light Actions in the Germination of Cocklebur Seeds: V. EFFECTS OF ETHYLENE, CARBON DIOXIDE AND OXYGEN ON GERMINATION IN RELATION TO LIGHT

Esashi, Y., Hase, S. and Kojima, K. 1987. Light actions in thegermination of cocklebur seeds. V. Effects of ethylene, carbondioxide and oxygen on germination in relation to light.–J.exp. Bot. 38: 702–710. Effects of ethylene, CO² and O² on the germination of after-ripenedupper cocklebur (Xanthium pennsylvanicum Wallr.) seeds wereexamined in relation to pre-irradiation by red (R) or far-red(FR) light In order to remove the pre-existing Pfr, seeds weresoaked in the dark for various periods prior to light irradiationand gas treatments. Regardless of light, 0.3 Pa C₂H₄ promotedgermination at 23 ?C, but it strongly inhibited germinationwhen applied at 33 ?C, the optimal temperature for the germinationof this seed. However, delayed application of C₂H₄ during 33?C incubation stimulated germination independently of lightin a similar manner to that seen at 23 ?C. It is, therefore,suggested that the germination-regulating action of C2H4 iscompletely independent of phytochrome. In contrast, the germination-promoting effect of 3–0 kPaCO₂ was pronounced only when the seeds were previously irradiatedby R, regardless of temperature, suggesting that CO₂ actionto promote germination depends upon Pfr. A synergism betweenCO₂ and C₂H₄ at 23 ?C was observed only in the germination ofseeds pre-irradiated by R, while at 33 ?C an antagonism occurredindependently of light. The stimulation of C₂H₄ production byCO₂ was most striking in the cotyledonary tissue pre-irradiatedby R. However, the R-dependent enhancement of CO₂-stimulatedC₂H₄ production was negated by the subsequent FR and it wasnot found in the presence of 1-aminocyclopropane-1-carboxylicacid (ACC). Moreover, the R dependency of the germination-promotingCO₂ effect disappeared in the presence of C₂H₄. The R-dependentC₂H₄ production enhanced by CO₂ may thus be involved, at leastpartially, in some step of conversion from methionine to ACC. The germination-promoting effect of C₂H₄, but not CO₂, was enhancedby O₂ enrichment regardless of light. However, the germination-promotingeffect of pure O₂ itself appeared to depend upon pre-irradiationwith R Key words: Carbon dioxide, cocklebur seed, ethylene, far-red light, germination, oxygen, red light, Xanthium pennsyloanicum     

葡萄属种子发育的物候、萌发行为及其对冷层积的反应

以3个起源和分布中心的不同葡萄种及其杂交种作为实验材料,观察了开花、果实和种子成熟的物候,研究了成熟种子的基本特性和最初萌发率以及冷层积对成熟种子萌发的影响。结果表明,葡萄种子具有休眠特性,生理成熟时都具有分化完全的胚,在休眠解除过程中胚的形态不发生变化;美洲种休眠最深,欧亚种休眠最浅,其它种类的葡萄种子的休眠程度介于美洲种和欧亚种之间;不同种葡萄花期、果实和种子成熟过程的物候存在差异,果实成熟与种子成熟不同步,其间隔时间越短,种子休眠程度越深。冷层积60天能有效地解除东亚种、欧亚种、欧山杂种和蘡欧杂种的种子休眠,但对解除美洲种、美洲种种间杂种和欧美杂种的种子休眠效果较差。