Studies in seed dormancy

Summary The dormancy of freshly harvested hazel seeds appears to be induced by inhibitors occuring mainly in the testa and pericarp. Although d abscisic acid may not be one of the natural inhibitors involved, d,l abscisic acid has been shown to strongly inhibit the germination of hazel seeds, probably through its antagonism towards the action of gibberellin. Dry storage of hazel nuts causes a deeper state of dormancy (secondary dormancy) to be superimposed on the primary dormancy. It is suggested that secondary dormancy consists of a block to gibberellin synthesis. The essential effect of chilling intact hazel seeds, which is the natural means of breaking their dormancy, may be to activate the mechanism for gibberellin synthesis, the subsequent synthesis of gibberellin being thought to occur at the germination temperature (20°C) and not at the chilling temperature (5°C).     

The Role of the Lens in Seed Imbibition and Seedling Vigour of Sesbania punicea (Cav.) Benth. (Leguminosae: Papilionoideae)

Seeds of Sesbania punicea (Cav.) Benth. are largely impermeable,and the natural, low, percentage which germinate are permeableat the lens. Scarifying seed in acid stimulates germinationand causes various types of damage to the testa, including thelens. Placing seeds in boiling water stimulates germinationby rendering the seeds permeable at the lens. Seeds were mechanicallyscarified at precise sites on the testa to assess influenceof the site and degree of damage on germination. Results indicatethat both the site of initial water entry and the rate of waterentry are important in determining seed and seedling vigour.Damaging the lens is more effective in stimulating germinationand reducing seedling abnormalities than complete excision ofthe lens. This effect can be reversed if seeds from which thelens was excised are allowed to imbibe gradually in inert osmotica.The leguminous lens thus acts as the site of initial water entryand regulates the rate of water uptake, thereby increasing seedand seedling vigour. Sesbania punicea (Cav.) Benth., hardseededness, lens, seed germination, imbibition damage, seedling vigour     

Hormonal Regulation of Seed Dormancy in Hazel (Corylus avellana L.) and Beech (Fagus sylvatica L.)

Dormancy in seeds of hazel (Corylus avellana L.) and beech (Fagussylvatica L.) has been studied with special reference to changesin growth-promoting and inhibiting substances during after-ripening.About 12 weeks at low temperature and under moist conditionsis necessary for complete after-ripening. Gibberellic acid,kinetin, and thiourea stimulate germination in dormant seedsbut have no effect on nuts with the pericarp intact. Gibberellin‘D’ is ten times more active than gibberellic acid.Bio-assays, following chromatographic fractionation of seedextracts, have revealed no significant changes in the concentrationsof auxins and inhibitors during after-ripening. Dwarf maize-leafsection assays have revealed low concentrations of gibberellin-likesubstances in purified extracts of chilled, dormant hazel seedbut no gibberellin activity in extracts of dormant seed. Gibberellinsare present in both dormant and germinating beech seeds butthere appear to be differences in the chromatographic patternof activity. The possible role of endogenous gibberellins inthe after-ripening process is discussed.     

Seed Dormancy in Acer pseudoplatanus L.: the Role of the Covering Structures

The present studies with Acer pseudoplatanus L. suggest thatthe covering structures play an important and multiple rolein the dormancy of the fruit. Whole fruits and seeds with thetesta intact required a period of chilling at 5 °C beforedormancy was broken whereas bare embryos germinated immediatelyat 20 °C without pretreatment. This suggested that dormancywas coat-imposed and that the testa was responsible for thiseffect. Germination of dormant seeds was inhibited by lightwhereas the non-dormant bare embryos showed little response.Studies on the manner in which the testa imposed dormancy onthe embryo indicated that restriction on oxygen uptake, wateruptake, mechanical restriction to embryo enlargement, and thepresence of germination inhibitors in the testa were not limitingfactors at this stage of dormancy. Results from leaching experimentssuggest that dormancy was the result of the restriction by thetesta of the outward diffusion of a germination inhibitor(s)present in the embryo. In seeds that had nearly completed theirstratification requirements, the covering structures seemedto act in a manner other than by preventing the leaching ofan inhibitor from the embryo. At this point the physical propertiesof the covering structures seem to determine any further delaysin germination by the mechanical restriction of embryo enlargementby the testa and by restriction of oxygen uptake by the pericarp.     

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     

Primary dormancy in tomato (Lycopersicon esculentum cv. Moneymaker): studies with the sitiens mutant

Intact wild-type tomato (Lycopersicon esculentum cv. Moneymaker)seeds do not complete germination to the same percentage orat the same speed as intact ABA-deficient sitiens (sit^w) mutantseeds when seeds of both genotypes are imbibed on polyethyleneglycol (PEG) solutions of –0.3 to –1.5 MPa osmoticpotential. However, if the thicker testas of wild-type seedsare removed (stripped) from the micropyle without damaging theendosperm, both the percentage and speed of germination at lowexternal water potential are similar to that of sit^w mutantseeds. Removing the micropylar end of the testa from sit^w seedsdid not enhance either the speed or percentage of germinationon PEG solution. Despite similar germination percentage and speed between strippedwild-type seeds and either stripped or intact sit^w seeds underosmotic stress, some differences in seed metabolism are evidentbetween genotypes. The activity of endo-ß-mannanasewas greater in the endosperm of sit^w mutant seeds compared tothe endosperm of wild-type seeds when seeds were exposed toosmotic stress. Although     

Effects of Priming and Endosperm Integrity on Seed Germination Rates of Tomato Genotypes: I. GERMINATION AT SUBOPTIMAL TEMPERATURE

The bases of differences in germination rates (GR_g = inverseof time to germination [t_g] of percentage g) among three cold/salt-toleranttomato (Lycopersicon esculentum Mill.) accessions (PI 341988,PI 120256, and PI 174263) and one cold/salt-sensitive tomatocultivar (T5) were investigated. The effects of seed priming(6 d imbibition in aerated –1.2 MPa polyethylene glycolsolution at 20 ?C followed by redrying) and of removing theendosperm/testa cap covering the radicle on the temperaturesensitivity of GR_g, and the interaction of these treatmentswith genotypes, were also examined. GR_g decreased linearly withdecreasing temperature for all genotypes and seed treatments.The minimum or base temperatures for germination (T_b) variedby 1 ?C among the tomato lines, so genotypic differences inGR_g were due to differing thermal time requirements for germination.The mean thermal time requirement for germination of T5 seeds was 22% and 19% greater than that of PI 341988 andPI 120256 seeds, respectively, but only 9% greater than thatof PI 174263 seeds. Seed priming did not lower Tb of any genotype,but significantly reduced by 24, 49, 41, and 49% in T5, PI 341988, PI 120256, and PI 174263, respectively,indicating that priming increased the rate at which the seedsprogressed towards germination when T>T_b, but did not lowerthe minimum temperature at which germination could occur. Primingincreased the GR_g of T5 seeds to equal or exceed those of control(non-primed) seeds of the cold/salt-tolerant genotypes at anyT>T_b, but the PI lines exhibited an even greater responseto priming. Times to germination within each seed lot were normallydistributed on a logarithmic scale. Priming increased the variancein tg within a seed lot when compared to control seeds. However,the variation in thermal time for germination between the 10thand 90th percentiles of the seed population (_T(10–90))was relatively unaffected by priming due to the reduction in in primed seeds. Removing the endosperm cap and testa opposite the radicle tip decreased almost 6-fold and and reduced T_b by 5 ?C in T5 and PI 341988,implicating processes in the endosperm/testa as the limitingfactors in germination at suboptimal temperatures. Key words: Lycopersicon esculentum Mill., tomato, genetic variation, seed priming, thermal time, germination rate     

Genetic Regulation of Seed Dormancy in Purshia tridentata(Rosaceae)

Seeds of Purshia tridentata, a shrub of semi-arid North America,require chilling to become non-dormant. Using seeds producedfrom controlled crosses in a common garden, we examined effectsof ovule parent, pollen parent, and year of production on germinationpercentage in response to chilling for 2 weeks at 2°C. Differencesamong ovule parents accounted for most of the variance in chillingresponse, and these differences (2 to 83% germination) wereconsistent across years. Differences among pollen parents werealso significant, producing a two- to five-fold difference inmean germination percentage. Differences among years were significantbut small. Ovule parent by year interactions showed that among-yearvariation in ripening environment did not affect ovule parentsequally. There was no significant pollen parent by year interaction,suggesting that the effect of maturation environment was mediatedthrough maternal tissues. In reciprocal crosses, two plantsthat showed contrasting dormancy levels as ovule parents producedseeds with similar dormancy as pollen parents, indicating thatthe genetic difference between them was at the testa level.Two plants that produced seeds with contrasting dormancy aspollen parents showed a similar but stronger pattern of contrastas ovule parents, showing that the genetic difference betweenthem was at both embryo and endosperm or testa level. Testaor endosperm genotype was primarily responsible for chillingresponse of intact seeds, while embryo genotype affected chillingresponse and also exercised primary control over low temperaturegermination rate, whether of excised embryos or of intact seeds.Copyright 2000 Annals of Botany Company Antelope bitterbrush, bitterbrush, common garden study, germination     

On the Nature of the Soaking Injury of Phaseolus vulgaris Seeds

Phaseolus vulgaris L. seeds are often killed or show signs ofsevere injury after they have been soaked in water for periodsexceeding a very few hours. But all ill effects can be largelyor completely prevented by any one of the following: (a) removingthe testa before or after soaking; (b) thorough or even partialdrying of the soaked seed; (c) draining the seed after cuttingoff its end portion; (d) treatment of the seed with hydrogenperoxide before, during, or after soaking. Air bubbled throughthe soaking water can aggravate the injury. By contrast, injuryfrom soaking can be prevented if oxygen has free access to theinterior of the seeds until they have imbibed about one-thirdof the water required for germination.It is suggested that,despite microbial complications at a later stage, the soakinginjury is caused at a critical early stage of germination bya deficient oxygen supply to the interior of the soaked seed,because during soaking the cavity between the cotyledons isflooded with an excess of water which remains trapped unlessforcibly removed.     

Seed and Pod Wall Development in Pisum sativum, L. in Relation to Extracted and Applied Hormones

Developing fruits of Pisum sativum, L., cv. ‘Alaska’,contain relatively large amounts of hormones, mainly concentratedin the embryos and liquid endosperm. A close relationship canbe demonstrated between changes in extractable amounts of gibberellins(mainly GA₂₀), auxins (methyl 4-chloroindol-3yl acetate andprobably 4-chloroindol-3yl acetic acid), and abscisic acid,and changes in growth rates of both the pod wall and seeds.Growth of the pod wall appears to depend largely on hormonessupplied by the seeds. Marked changes in the germination capacity of the maturing seedsare closely associated with changes in extractable amounts ofmethyl-4-chloroindol-3yl acetate and abscisic acid. It is believedthat high concentrations of these substances in the embryo,rather than any restriction imposed by the testa, may preventprecocious germination of the seeds     

Starch Synthesis During Dormancy Breakage in Oilseed of Corylus avellana

Dormancy in seed of Corylus avellana L. (hazel) is broken bya sustained period of cold stratification. During this timeboth cytological and metabolic changes occur. Starch was presentinitially at a low level but increased by 20% in the embryonicaxes of hazel seeds during stratification at 5 °C, whileit decreased rapidly and then remained constant in the embryonicaxes from seeds held at 20 °C. Histochemical study confirmedthis analytical result. A comparison of the developmental patternof starch level with bound and soluble ADP glucose-starch synthaseactivity at 5 and 20 °C showed that the accumulation ofstarch in the embryonic axes followed an enhanced activity ofthe granule-bound ADP glucose-starch synthase. Cold stratificationresulted in an increase in starch content, which was probablyas a result of gluconeogenesis from products of reserve lipidhydrolysis. Corylus avellana L., hazel, starch, ADP glucose-starch synthase, stratification     

The Role of ABA in the Control of Apple Seed Dormancy Re-appraised by Combined Gas Chromatography-Mass Spectrometry

Measured by GC—MS²—SIR³, endogenous ABA⁴ in embryonicaxes of seeds of Malus pumila L. cv. Golden Delicious decreased8-fold and cotyledon ABA by only 60%, during 10–50 d ofstratification at 5 ?C, after ABA leaching during an initial24 h soaking. During stratification, the percentage germinationof embryos transferred to 17?C showed a significant linear dependenceon log_e of ABA levels in the axes at transfer. Between 50 and70 d, ABA levels increased markedly in axes and testa both ofstratified seeds and seeds allowed to re-dry at 17 ?C afterinitial soaking. The ability of fully stratified axes with elevatedendogenous ABA to germinate indicated that stratification haddecreased their ABA sensitivity. Changes in cotyledon ABA couldnot account for the promotory effect of cotyledons on germinationduring the first 30 d of stratification. Loss of testa inhibitionof germination during stratification was not linked with changesin testa ABA. Stratification markedly increased the sequestrationin the axes of exogenous ABA supplied via the cotyledons. Changesboth in axis ABA levels and sensitivity were thus correlatedwith dormancy release, but subject to modifying control by thecotyledons and testa not involving ABA. Rehydration of driedseeds affected axis ABA later during storage via mechanismsunconnected with dormancy. Key words: ABA, seed dormancy, stratification     

新疆干旱区植物藜的种子异型性及其萌发机理

新疆干旱区分布的植物藜(Chenopodium album)的种子有黑色和褐色两种类型。对藜的异型性种子从形态结构、不同环境因素及激素或化学物质对萌发的影响以及同工酶谱等方面进行了研究,并对其萌发及适应异质环境的机理进行了讨论。结果表明:(1)藜的异型性种子在形态结构、萌发休眠特性等方面都存在明显差异:黑色种子种皮厚且硬,休眠,萌发慢,萌发率低;褐色种子种皮薄而软,不休眠,萌发快且萌发率高;(2)黑色种子的休眠可通过切除胚根外缘种皮得以完全解除;(3)赤霉素、乙烯利对黑色种子的萌发无明显促进作用;KNO3可较显著促进黑色种子的萌发;协同使用乙烯利和KNO3时,可显著提高黑种子萌发率,完全打破休眠;(4)黑色种子和褐色种子的酯酶、过氧化物酶及过氧化氢酶同工酶谱带存在差异;(5)黑色种子的萌发需要光照,而褐色种子则对光不敏感;低温贮藏对二者的萌发均无显著影响,尽管黑色种子的萌发率有波动。研究结果初步显示黑色种子的休眠是内源(胚)和外源(种皮)因素共同所致。藜的种子异型性及其萌发机理的形成是其对新疆干旱区异质化环境的高度适应。     

Removal of the pericarp and testa of seeds of japonica and indica rice (Oryza sativa) at various oxygen concentrations has opposite effects on germination

The germination of intact, dehusked, and peeled seeds (caryopses) of the japonica rice cultivar Sasanishiki, harvested 30, 40, 47 and 60 days after anthesis, and of the indica rice cultivar Assam IV, harvested 14 and 28 days after anthesis, was examined. Dehusking strongly inhibited germination of Sasanishiki seeds, with the exception that seeds harvested 30 days after anthesis gave minimal germination percentages even when left intact. Peeling (removal of the pericarp and testa) restored or enhanced germination, and 60–100% of seeds germinated after 10 days. By contrast, the rank order of germination of Assam IV seeds was intact, dehusked, and peeled seeds, with peeled seeds yielding germination percentages of 100%. In Sasanishiki, inhibition of germination of peeled seeds was observed at reduced oxygen concentrations (1–4% oxygen). This inhibition might explain the inhibitory effects of dehusking on germination of seeds from the japonica cultivar. It is possible that the testa and pericarp, which cover the embryos of dehusked seeds, acted as a barrier to the diffusion of oxygen to the embryo.     

The Effect of Temperature on the Supply of Oxygen to Embryos ofIntact Acer pseudoplatanus L. Seeds

The effect of temperature on various factors affecting O₂ supplyto enclosed embryos of A. pseudoplatanus L. seeds was investigatedusing a simple model for O₂ consumption by intact seeds. Lowtemperatures which induced germination had little effect onthe diffusivity of the testa although an increase in diffusivityoccurred immediately prior to radicle emergence. The O₂ concentrationsurrounding an enclosed embryo was found to be 5–10% thatof the solution bathing the seeds at storage temperatures. Therespiratory capacity of the embryos was found to be stronglydependent on temperature and, largely as a result of this, atchilling temperatures the O₂ concentration surrounding the enclosedembryo approached that in the ambient solution. The apparentArrhenius activation energy for O₂ consumption was considerablygreater below 17 °C than above this temperature. Althoughinsufficient evidence in themselves for a relationship betweenO₂ availability and seed germination, the results do provideevidence for a mechanism by which such a relationship may beassociated with seeds requiring chilling to induce germination.     

Changes in Phenolic Inhibitors in Seeds of Acer saccharum during Stratification

Phenolics in sugar maple seeds were identified by colour reactions,u.v. spectroscopy and gas-liquid chromatography. Changes intheir inhibitory activity during stratification were also followedusing several bioassays. p-Coumaric, o-coumaric, and ferulicacids were found in all tissues. The highest concentrationswere noted in the embryonic axis and testa, but total amountsper fruit were highest in cotyledons and pericarp. p-Coumaricacid was the principal phenolic in dry seeds, and it declinedsubstantially in all tissues during the first half of stratificationwith little subsequent change thereafter. Decreases in o-coumaricand ferulic acids per seed were small by comparison, neverthelesslosses ranged between 36 and 68% of the original concentrations.Bioassays confirmed that the three endogenous phenolics possessedmarked inhibitory properties that diminished with the progressof stratilication, but only the compound identified as p-coumaricacid showed major inhibition in seed germination tests.     

The Influence of Light and Potassium Nitrate on the Dormancy and Germination of Avena fatua L. (wild oat) Seed Stored Buried Under Natural Conditions

The influence of light and potassium nitrate on the dormancyand germination of Avena fatua L. seed stored buried under naturalconditions was studied over a 13 month period. Natural emergenceof seedlings from buried seed in relation to rainfall and temperatureregimes and fluctuation was also monitored. The present findingsshow that potassium nitrate is stimulatory to germination onlyin the light and, further, that seeds from two positions inthe spikelet respond differently. Key words: Dormancy, Light, Nitrate     

PHOTOCONTROL AND TEMPERATURE DEPENDENCE OF GERMINATION OF RUMEX SEEDS

1. Light is not obligatory for the germination of the seed ofRumex obtusifolius L. subsp.agrestis DANSER, which has beenregarded as being a typical light sensitive seed. Even in continuousdarkness, a short period of high (30°) or low temperature(5°) treatment evokes germination very readily. 2. Germination is markedly promoted by 1 min exposure to a redlight and this red light effect is completely removed by 1-hrexposure to a far-red light. Alternations of the red and far-redradiation bring about an alternate promotion and inhihibitionof germination. 3. When a dark interval is inserted between the red and thefar-red treatments, inhibition of germination becomes less distinctas the duration of darkness increases. When the seeds are irradiatedwith far-red prior to red, with an inserted darkness, germinationpromotion due to the red light also decreases with the durationof inserted darkness. 4. Complicated interdependence between the light and temperatureeffects are demonstrated. This suggests a participation of somereactants besides pigments in the photoreaction. 5. The observed interdependence between the light and temperatureeffects on the germination of Rumex seeds implies that, if,as BORTHWICK has assumed, two forms of pigment, viz., a far-red-absorbingform and a red-absorbing one, are participating in the photoreaction,they should be presumed to coexist from the start of imbibition. (Received September 27, 1960; )     

Fusarium oxysporumf. sp.orthoceras, a Potential Mycoherbicide, Parasitizes Seeds ofOrobanche cumana(Sunflower Broomrape): a Cytological Study

Fusarium oxysporumf. sp.orthoceras, a potential agent for biologicalcontrol of the root-parasitic weedOrobanche cumanain sunflower,attacks underground developmental stages, such as shoots, tuberclesand seed germ tubes. The germination of inoculated seed wassignificantly reduced, and cytological investigation showedthatF. oxysporumhad penetrated and colonized dormant seeds ofO.cumana. Germ tubes of microconidia penetrated all parts of thethick, complex seed testa, and seed contents were completelydestroyed. Hyphae dissolved the endosperm cell walls and metabolizedthe cytoplasm which was rich in lipid- and protein-bodies. Thus,the inoculation of soils withF. oxysporummay be used to reducetheOrobancheseed bank.Copyright 1999 Annals of Botany Company Fusarium oxysporum, sunflower broomrape,Orobanche cumana, seeds, electron microscopy, ultrastructure, infection process, biological control, bioherbicide, mycoherbicide.