Control of seed coat thickness and permeability in soybean : a possible adaptation to stress

Although the seed coat, through its thickness and permeability, often regulates seed germination, very little is known about the control of its development. Using soybean (Glycine max [L.] Merrill) explants, podbearing cuttings in which defined solutions can be substituted for the roots, we have demonstrated that cytokinin and mineral nutrients moving through the xylem can control soybean seed coat development. Lack of cytokinin and minerals in the culture solution, causes a thicker, less permeable seed coat to develop. The seeds with thickened coats will imbibe water rapidly if scarified; furthermore, these scratched seeds also germinate and produce normal plants. Inasmuch as stress (e.g. drought) decreases mineral assimilation and cytokinin production by the roots, the resulting delay in germination could be an adaptive response to stress.     

Imbibition phases and germination response of Mimosa bimucronata (Fabaceae: Mimosoideae) to water submersion

Mimosa bimucronata is a pioneering tree that occurs predominantly in moist lowlands, floodplains and on margins of rivers and lakes in Latin America. The effect of submergence on seed germination in M. bimucronata was firstly studied. Patterns of water absorption by M. bimucronata seeds were investigated thereafter to assess the imbibition phases of scarified and unscarified seeds. The germination percentage was significantly higher in scarified than in unscarified seeds, and the velocity of seed germination also increased considerably in scarified seeds. Submergence duration did not significantly affect germination percentages of scarified and unscarified seeds. Therefore, seed viability after submersion suggests that M. bimucronata may display hydrochorous dispersal and also that seeds are able to germinate successfully in areas with frequent seasonal flooding. With respect to imbibition phases, phase II was very short or even absent for scarified and unscarified seeds; therefore, a plateau, where water absorption by seeds is established, was not observed. Finally, we verified that the passage from phase I to III was very tenuous and took a long time in seeds without scarification.     

Seed dormancy of Cardiospermum halicacabum (Sapindaceae) from three precipitation zones in Sri Lanka

• This study investigated seed germination of Cardiospermum halicacabum, a medicinally important invasive species.
• We compared mass, moisture content (MC), dormancy and dormancy‐breaking treatments and imbibition and germination of scarified and non‐scarified seeds of C. halicacabum from a low‐elevation dry zone (DZ), low‐elevation wet zone (WZ1) and mid‐elevation wet zone (WZ2) in Sri Lanka to test the hypothesis that the percentage of seeds with water‐impermeable seed coats (physical dormancy, PY) decreases with increased precipitation.
• Seed mass was higher in WZ2 than in DZ and WZ1, while seed MC did not vary among the zones. All scarified DZ, WZ1 and WZ2 and non‐scarified DZ and WZ1 seeds imbibed water, but only a few non‐scarified WZ2 seeds did so. When DZ and WZ1 seeds were desiccated, MC and percentage imbibition decreased, showing that these seeds have the ability to develop PY. GA₃ promoted germination of embryos excised from fresh DZ and WZ1 seeds and of scarified WZ2 seeds.
• At maturity, seeds from DZ and WZ1 had only physiological dormancy (PD), while those from WZ2 had combinational dormancy (PY+PD). Thus, our hypothesis was not supported. Since a high percentage of excised embryos developed into normal seedlings; this is a low‐cost method to produce C. halicacabum plants for medicinal and ornamental purposes.

     

Germination requirements inCarex kobomugi (Sea Isle)

Laboratory and field germination experiments inCarex kobomugi seeds were pursued to clarify their germination requirements and availability of the requirements in the field. In the laboratory experiments, more than 50% of the seeds ofC. kobomugi germinated under 35/30C or 25/20C when they were scarified with 98% H₂SO₄ after removal of their utricles, and chilled in moist condition for 28 to 42 d. Seeds with utricles or those without scarified with H₂SO₄ did not germinate. Seeds sown at 10-cm depth at the Kado-ori coast on 11 February 1991 after soaked in H₂SO₄ showed 40% germination by 29 April 1991, whereas those without H₂SO₄ treatment did not germinate. These results suggest that seed-coat impermeability and embryo immaturity are possible causes of the dormant state in seeds ofC. kobomugi ripen in summer. In the field, the moist-chilling condition is available in winter and the seeds can germinate in the following spring if the seed-coat impermeability is relaxed before winter.     

Uptake of Gibberellic Acid in Intact and Scarified Seeds of Barbarea vulgaris

Uptake of gibberellic acid as a function of duration of exposure, external concentration, and seed lot was measured in seeds of yellow rocket (Barbarea vulgaris R. Br.) by means of lettuce hypocotyl bioassays of the acidic, basic, or neutral fractions of seed extracts and by uptake of ¹⁴C-GA₃. In intact seeds, where mM levels of GA₃ promoted only 25% germination, uptake was completed within 24 h of exposure. The maximum uptake was about 0.2% of external amount. Although germination promotion by GA₃ differed among seedlots of yellow rocket, relative uptake (percentage of the external GA₃) was nearly the same. The relative rate of uptake of GA₃ was similar for scarified and intact seeds, but germination was promoted in scarified seeds by much lower levels of GA₃ than in intact seeds. Total uptake in scarified seeds was much higher, however (about 10% of the external amount). In seeds imbibed in H₂O, practically no endogenous GA-like activity was detected in either the acidic, basic, or neutral fractions. It was also apparent that intact seeds could take up quantities of GA₃ that failed to promote germination, but were comparable to quantities that promoted germination in scarified seeds.     

Influence of factors affecting germination on respiration of Phacelia tanacetifolia seeds

Summary Germination of the seeds of Phacelia tanacetifolia is inhibited by light. Removal of that part of the covering structures of the seeds which directly covers the radicle allows full germination in light. The rate of O₂ uptake in the seeds increases following imbibition, and reaches the same steady rate in light and in darkness after 3 hr. From the 14 th hour on, dark-imbibed seeds show a linear increase in the rate of respiration. This increase is not observed in dormant seeds incubated in light. In normal dark germination, protrusion of the radicle begins at 12 th hour following soaking, and by the end of 18 th hour approximately 60% of the seeds have germinated. The seeds which have been scarified at the radicle end and germinate readily in light show a steady increase in Q _{O 2}. If scarified seeds are allowed to imbibe 0.3 M mannitol and are then incubated in light, the embryo does not grow and the pattern of O₂ uptake becomes identical with that of intact seeds in light. Mannitol, however, does not inhibit respiration by itself. These observations indicate that the increased O₂ uptake is the result rather than the cause of seed germination, and that light does not cause dormancy by inhibiting O₂ uptake. Measures effective in releasing dormancy (dark incubation, mechanical scarification, gibberellin treatment) do not induce germination by facilitating oxygen entry.This work was supported in part by the Jane Coffin Childs Memorial Fund for Medical Resarch, and by the U. S. Atomic Energy Commission under Contract No. AT (11-1)-1338.     

The Effects of Gibberellic Acid and Scarification on the Seed Dormancy and Germination in Luzula spicata

Luzula spicata L. seeds are completely dormant at maturity. A germination inhibitor is present at the micropylar end. Normally, the only effective means of eliciting germination is a precise scarification of the micropylar end which inactivates the inhibitor. Exogenous application of gibberellic acid, kinetin, KNO₃, and thiourea have no affect on the dormancy of unscarified seeds. Scarification of the hilar end of the seed does not elicit germination, but when gibberellic acid is applied to the hilar scarified seeds moderate germination results. Presumably, these seeds are dormant due to a deficit of endogenous gibberellin; a condition which can be overcome by the application of gibberellic acid to seeds scarified at a site in itself ineffective in producing germination. Apparently the gibberellic acid serves to initiate amylase activity in the endosperm, overcoming the inhibitor block. Luzula spicata seed dormancy is apparently unique in that a germination inhibitor is operative in conjunction with the commonly recognized gibberellin-amylase mechanism.     

Bet‐hedging and germination in the Australian arid zone shrub Acacia ligulata

The diaspore of the Australian arid zone shrub Acacia ligulata is dispersed by birds and ants. To investigate the benefits of providing a dispersal structure attractive to both groups, we compared the germination response and viability of seeds eaten by birds, handled by ants or collected from trees to simulated precursors of germination: scarification, fire and rainfall were simulated. Seed germination and viability were related to the degree of preheating disturbance to the seed coat. Heating increased the germinability of seeds not scarified or eaten by birds. In the absence of heating, ingestion by birds increased germinability. Heating increased the mortality of seeds. Our results suggest that ingestion of seeds by birds may break seed dormancy and hence enable some seeds to germinate soon after dispersal. Alternatively, seeds not eaten by birds are likely to remain dormant until sufficiently scarified by soil or stimulated by fire. Consequently, in areas such as the Simpson Desert, A. ligulata may be able to use a range of seedling establishment ‘windows’ provided by monsoon rains, post‐fire environments and unseasonal winter rains, and also spread the risk of unsuccessful seedling establishment by retaining dormant seeds in the seedbank.     

PHYSIOLOGY OF LIGHT-REQUIRING GERMINATION IN ERAGROSTIS SEEDS

The photorequirement for the germination of Eragrostis seedsdecreases with the progress of their after-ripening, and thegermination occurs whether in continuous light or in darknessat the final stage of after-ripening. The dehydration of seedsor the puncturing of seed coats also results in a decrease ofphotorequirement for germination. The rate of water absorptionof seeds increases with the germination capacity under continuousdark condition. However, there is no correlation between therespiration rate and the germination capacity; respiration isstimulated in the punctured seeds, but not in the after-ripenedseeds. The after-ripened or punctured seeds which no longer have aphotorequirement become light-sensitive again, when they areallowed to germinate in the air of low oxygen concentrations.The assumption is presented that the permeability to oxygenof the seed coat may be a factor controlling the seed germinationof this species. (Received August 21, 1964; )     

Sensitivity cycling in physically dormant seeds of the Neotropical tree Senna multijuga (Fabaceae)

• Cycling of sensitivity to physical dormancy (PY) break has been documented in herbaceous species. However, it has not been reported in tree seeds, nor has the effect of seed size on sensitivity to PY‐breaking been evaluated in any species. Thus, the aims of this study were to investigate how PY is broken in seeds of the tropical legume tree Senna multijuga, if seeds exhibit sensitivity cycling and if seed size affects induction into sensitivity.
• Dormancy and germination were evaluated in intact and scarified seeds from two collections of S. multijuga. The effects of temperature, moisture and seed size on induction of sensitivity to dormancy‐breaking were assessed, and seasonal changes in germination and persistence of buried seeds were determined. Reversal of sensitivity was also investigated.
• Fresh seeds were insensitive to dormancy break at wet–high temperatures, and an increase in sensitivity occurred in buried seeds after they experienced low temperatures during winter (dry season). Temperatures ≤20 °C increased sensitivity, whereas temperatures ≥30 °C decreased it regardless of moisture conditions. Dormancy was broken in sensitive seeds by incubating them at 35 °C. Sensitivity could be reversed, and large seeds were more sensitive than small seeds to sensitivity induction.
• Seeds of S. multijuga exhibit sensitivity cycling to PY‐breaking. Seeds become sensitive during winter and can germinate with the onset of the spring–summer rainy season in Brazil. Small seeds are slower to become sensitive than large ones, and this may be a mechanism by which germination is spread over time. Sensitive seeds that fail to germinate become insensitive during exposure to drought during summer. This is the first report of sensitivity cycling in a tree species.

     

Phytochrome et germination des semences de Rumex

The Rumex alpinus L. achenes show a special type of positive photosensitivity: several red light irradiations are necessary to induce optimal germination. The achenes deprived of their pericarp lose their photosensitivity and germinate readily in the dark. By spectrophotometry in vivo, the presence of the Pr form of phytochrome is revealed in these seeds. Short periods at extreme temperatures (30°C or 5°C) induce a good proportion of achenes to germinate. The gibberellins are inefficient on achene germination contrary to benzyladenine which exhibits some activity. A slight improvement of GA₄ effects has been detected on scarified fruits.     

Development of seed coat-imposed dormancy during seed maturation in Cynoglossum officinale

The relationship between seed phenolics and appearance of seed coat–imposed dormancy during seed development in Cynoglossum officinale L. was studied. Up to 24 days after anthesis, seeds failed to germinate upon imbibition in Petri dishes at 25°C. At 44 days after anthesis, seeds were fully germinable; removal of seed coats did not improve their germination or O₂ uptake. At 72 days after anthesis, mature seeds at the base of the cyme did not germinate unless their coats were removed. Removal of seed coat also stimulated O₂ uptake at this harvest date. The methanol-soluble phenolic content of the seeds increased during the early stages of seed development, in both the seed coat and the embryo. As seed development continued, the methanol-soluble phenolic content of the embryo stabilized, but that of the seed coat declined. This decline was associated with an increase in the thioglycolic acid–soluble phenolics, presumably lignins, in the seed coat. These results suggest that polymerization of methanol–soluble phenolics into lignins in the seed coat during later stages of seed development renders the seed coat of C. officinale impermeable to 03, and thus keeps the seed dormant.     

Induction of Secondary Dormancy in Chenopodium bonus-henricus L. Seeds by Osmotic and High Temperature Treatments and Its Prevention by Light and Growth Regulators

Factors controlling the establishment and removal of secondary dormancy in Chenopodium bonus-henricus L. seeds were investigated. Unchilled seeds required light for germination. A moist-chilling treatment at 4 C for 28 to 30 days removed this primary dormancy. Chilled seeds now germinated in the dark. When chilled seeds were held in the dark in −8.6 bars polyethylene glycol 6000 solution at 15 C or in water at 29 C a secondary dormancy was induced which increased progressively with time as determined by subsequent germination. These seeds now failed to germinate under the condition (darkness) which previously allowed their germination. Continuous light or daily brief red light irradiations during prolonged imbibition in polyethylene glycol solution at 15 C or in water at 29 C prevented the establishment of the secondary dormancy and caused an advancement of subsequent germination. Far red irradiations immediately following red irradiation reestablished the secondary dormancy indicating phytochrome participation in “pregerminative” processes. The growth regulator combination, kinetin + ethephon + gibberellin A₄+A₇ (GA₄₊₇), and to a relatively lesser extent GA₄₊₇, was effective in preventing the establishment of the secondary dormancy and in advancing the germination or emergence time. Following the establishment of the secondary dormancy by osmotic or high temperature treatments the regulator combination was relatively more active than light or GA₄₊₇ in removing the dormancy. Prolonged dark treatment at 29 C seemed to induce changes that were partially independent of light or GA₄₊₇ control. The data presented here indicate that changes during germination preventing dark treatment determine whether the seed will germinate, show an advancement effect, or will become secondarily dormant. These changes appear to be modulated by light and hormones.     

FACTORS INFLUENCING SEED GERMINATION IN SALICORNIA PACIFICA VAR. UTAHENSIS

The halophyte, Salicornia pacifica var. utahensis (Tiderstorm) Munz produces seed under high salinity conditions, and deposits its seed on saline soil. Experiments were conducted to determine the effect of salinity, temperature and growth regulators on germination. Results indicate that the seeds can germinate at very high salt concentration (5% NaCl). Germination was sensitive to the changes in temperature regimes. At higher 30–20 C, light-dark sequence, no germination occurred at 3, 4 and 5% NaCl treatments. On the other hand, 30% germination did occur at 5% NaCl treatment at a temperature regime of 15–5 C. These seeds required light for germination. Only 50% germination occurred in the non-saline control in the dark and the addition of NaCl further reduced germination. The GA₃ partially alleviated the inhibitory effect of NaCl and darkness. Kinetin did not promote germination.     

Effects of seed burial on germination,protein mobilisation and seedling survival in Dodonaea viscosa

Ecological restoration of disturbed areas requires substantial knowledge of the germination of native plants and the creation of novel methods to increase seedling establishment in the field. We studied the effects of soil matrix priming on the germination of Dodonaea viscosa seeds, which exhibit physical dormancy. To this end, we buried both pre‐scarified (in H₂SO₄, 3 min) and non‐pre‐scarified seeds in the Parque Ecológico de la Ciudad de México. After seeds were unearthed, they were post‐scarified for 0, 2, 6 and 10 min and their germination percentages compared to the germination of a control batch of laboratory‐stored seeds. For both control and unearthed seeds, the protein pattern was determined in the enriched storage protein fraction in SDS‐PAGE gels stained with Coomassie blue. Percentage germination increased as the scarification time increased. Pre‐scarification significantly increased percentage germination of post‐scarified seeds in relation to the control and non‐pre‐scarified seeds. In seeds unearthed from the forest site, the buried pre‐scarified seeds had relatively high percentage germination, even in the absence of post‐scarification treatment. A 48‐kDa protein was not found in unearthed, pre‐scarified seeds nor in the control germinated seeds, indicating that mobilisation of this protein occurred during soil priming. Burying seeds for a short period, including the beginning of the rainy season, promoted natural priming, which increased protein mobilisation. Functionally, priming effects were reflected in high percentage seedling survival in both the shade house and the field. Seed burial also reduced the requirement for acidic post‐scarification.     

Seeds germination of Caesalpinia paraguariensis (Fabaceae): scarificator agents and cattle effects]

The tree Caesalpinia paraguariensis grows in the Chaco region, Argentina. Fruits are indehiscent with many seeds. This species is an important source of wood and the fruits are consumed by cattle in Salta province. We studied seed germination under chemical, mechanical and biological scarification. Seeds from controls (without scarification) and those with biological scarification had a smaller (and similar) germination rate. The non-germinated seeds from biological treatments were mechanically scarified and their germination rate was similar to others under the mechanical treatment. Passage by digestive tracts would not enhance germination because viable seeds are still dormant due to their hard coats.     

VARIATION IN SEED WEIGHT AND ITS EFFECTS ON GERMINATION IN PASTINACA SATIVA L. (UMBELLIFERAE)

Pastinaca sativa (wild parsnip) produces seeds on the primary, secondary, and tertiary umbels of the flowering stalk. Within plants, variation in seed weight is about twofold. Secondary and tertiary seed weight is 73% and 50% of primary seed weight, respectively. Maximum variation in seed weight between plants is sixfold when tertiary seeds from a small plant are compared to primary seeds from a large plant. Within an umbel order, variation in seed weight between plants is correlated with plant size. Under autumn germinating conditions in the laboratory, final germination of seeds from different umbel orders does not differ but smaller seeds germinate more rapidly than larger seeds. Under spring germination conditions in the laboratory, significantly more primary and secondary seeds germinate than tertiary seeds and the rate of germination is independent of seed weight. Field germination of seeds from different umbel orders produces similar results except that in the spring both secondary and tertiary seed germination is lower than that of primary seeds. These results suggest that with respect to seed germination characteristics small seeds may have a competitive advantage over large seeds in the autumn because they germinate more quickly, but in the spring small seeds are at a disadvantage because they have lower overall germination. Because most germination in the field occurs in the spring, population recruitment from small seeds is likely to be substanially less than that from large seeds.     

Effects of various pretreatments on seed germination of<Emphasis Type="Italic">Calystegia soldanella</Emphasis> (Convolvulaceae), a coastal sand dune plant

We tested various pretreatments to enhance the germination of ‘Sea bells’ (Calystegia soldanella Roem. et Schult). One experiment, which improved the germination rate by 70%, involved first scarifying the seed coats, then immersing the seeds in 50 ppm GA₃ for 24 h. A GA₃-alone pretreatment did not increase germination. However, the most effective method, with a 100% success rate, included 3 h of acid pretreatment with 98% H₂SO₄. We also used scanning electron microscopy (SEM) to examine the seed-coat surfaces and cross-sections of dry seeds. SEM showed structural differences between seeds that were not treated and those exposed to 98% H₂SO₄. The latter treatment allowed the seed coats to crack and break, thereby disrupting their physical dormancy.     

Influence of the Seed Envelope and Growth Regulators Upon Seed Dormancy in Witchweed (Striga lutea Lour.)

Seed of the angiospermous parasite, witchweed (Striga luteaLour.), normally germinate poorly or not at all unless adequatelypretreated and exposed to a germination stimulant which is obtainedfrom plant root exudates. Under certain conditions, scarificationof the seed envelope promoted germination in the absence ofthe stimulant. With adequately (2-week) pretreated seed, a cutor puncture through the aleurone at the radicular end inducedtypical germination. A puncture through the aleurone elsewhereon the seed induced little or no germination. A cut throughthe aleurone at the cotyledon end or centre of the seed inducedatypical germination. A puncture through the aleurone elsewhereon the seed induced little or no germination. A cut throughthe aleurone at the cotyledon end or centre of the seed inducedatypical germination in which the radicle elongated but didnot penetrate the intact envelope over the radicle. Incubationin oxygen did not promote germination regardless of the siteof seed scarification. Ten per cent carbon dioxide reduced thegermination of punctured seed. Seed germinated equally wellwhen scarified over the radicle and incubated in air, nitrogen,darkness, or light. Brief treatments with sulphuric acid alsoinduced germination. The softening effect of stimulatory acidtreatments upon the aleurone was first evident 6 h later andonly occurred at the radicular end of the seed. Indol-3yl-aceticacid inhibited and ethylene stimulated germination of scarifiedand non-sacrified seed. Gibberellic acid(GA₃) had no apparenteffect upon nonscarified seed but promoted germination of scarifiedseed. Inadequately (I-day) or excessively (12- to 16-week) pretreatedseed germinated poorly or not at all when treated with the germinationstimulant, ethylene, GA₃, or sulphuric acid. Some seed germinatedslowly when scarified over the radicle, but the germinationrates and totals were less than those of scarified seed pretreatedfor 2 weeks. Additions of stimulant, ethylene or GA₃, aceleratedthe germination rates of scarified but inadequately pretreatedseed. Of the treatments tested, only GA₃, increased the slowgermination of the excessively pretreated scarified seed. Results indicated that the aleurone restrained radicle elongation.Scarification over the radicle removed the restraint and permittedradicle emergence. However, the ability of the radicle to elongate,as influenced by time of seed pretreatment and exogenous stimulant,GA₃ or ethylene, determined whether or not the scarified seedgerminated.     

Combinational dormancy in seeds of Sicyos angulatus (Cucurbitaceae,tribe Sicyeae)

Seeds with a water‐impermeable seed coat and a physiologically dormant embryo are classified as having combinational dormancy. Seeds of Sicyos angulatus (burcucumber) have been clearly shown to have a water‐impermeable seed coat (physical dormancy [PY]). The primary aim of the present study was to confirm (or not) that physiological dormancy (PD) is also present in seeds of S. angulatus. The highest germination of scarified fresh (38%) and 3‐month dry‐stored (36%) seeds occurred at 35/20°C. The rate (speed) of germination was faster in scarified dry‐stored seeds than in scarified fresh seeds. Removal of the seed coat, but leaving the membrane surrounding the embryo intact, increased germination of both fresh and dry‐stored seeds to > 85% at 35/20°C. Germination (80–100%) of excised embryos (both seed coat and membrane removed) occurred at 15/6, 25/15 and 35/20°C and reached 95–100% after 4 days of incubation at 25/15 and 35/20°C. Dry storage (after‐ripening) caused an increase in the germination percentage of scarified and of decoated seeds at 25/15°C and in both germination percentage and rate of excised embryos at 15/6°C. Eight weeks of cold stratification resulted in a significant increase in the germination of scarified seeds at 25/15 and 35/20°C and of decoated seeds at 15/6 and 25/15°C. Based on the results of our study and on information reported in the literature, we conclude that seeds of S. angulatus not only have PY, but also non‐deep PD, that is, combinational dormancy (PY + PD).