Caryopsis germination and seedling emergence in an inland dune dominant grass Leymus secalinus

Leymus secalinus (Georg.) Tzvel. (Poaceae) is a dominant sand dune grass inhabiting the Mu-Us Sandland, semiarid China. Freshly harvested caryopses (seeds) are in non-deep physiological dormancy (non-deep PD) because of low percentage and slow rate of germination. Experiments were conducted to examine the effects of temperature, cold stratification, caryopsis coat scarification or partial removal of endosperm and sand burial on caryopsis dormancy, germination and seedling emergence. Caryopsis germination was significantly influenced by duration of cold stratification, temperature and their interactions. After 8 weeks of cold stratification, caryopsis germination percentage at 30 °C reached to 90%, equally in light or darkness. Rate and percentages of germination were also hastened and increased by scarifying the caryopsis coat or by artificial removal of different proportions of the endosperm. However, seedling developmental characteristics were significantly influenced by the proportion of the endosperm that remained in the caryopses. Seedling emergence, caryopsis germination and enforced dormancy in sand were significantly affected by sand burial depth. As sand burial depth increased, caryopsis germination and seedling emergence decreased whereas caryopsis enforced dormancy increased. 1–2 cm was the optimal depths for caryopses germination and seedling emergence. Although there were still 30% caryopses germinated at 8 cm, the maximal burial depth for seedling emergence was only 4 cm. The partial germination strategy regulated by non-deep PD, temperature and sand burial ensures that only a few caryopses germinated each time and may reduce risks for seedling survival.     

Seed dormancy in Avena fatua: Interacting effects of nitrate, water and seed coat injury

In experiments conducted under controlled conditions. KNO₃ (50 or 100 m M ) promoted germination of a dormant strain (AN 474) of Avena fatua when either one or two holes were pierced in the lower (adaxial) surface of the caryopsis in contact with the nitrate solution. Germination was increased by increasing either the KNO₃ concentration or the number of holes in the seed coat. The germination response induced by the application of water to a hole pierced in the upper surface of the caryopsis was. increased by pre-treatment of the intact caryopsis with KNO₃. Treatment with either 50 or 100 m M KNO₃ caused a transient reduction in embryo water content of intact cary-opses, but increased the nitrate and amino- N content of pierced caryopses prior to germination. Supplying a 100 m M solution of KNO₃ to pierced caryopses reduced the total water potential and osmotic potential of the embryo, and increased its pressure potential by the same amount as an equimolar solution of KC1; however, while both treatments promoted germination, the KNO₃ induced more rapid germination than the KCI. Both treatments also increased the K⁺ content of the embryo, the KNO₃ again having the greater effect. These results are consistent with the hypothesis, based on our previous investigations, that KNO₃ promotes germination of dormant caryopses by accumulating in the embryo where it acts osmotically to increase water uptake. It is also postulated, that, in contrast to KCI, KNG₃ may combine an osmotic effect on water uptake with a nutritional effect on protein synthesis.     

Effect of fusicoccin and gibberellic acid on primary dormant Avena fatua caryopses

Fusicoccin induced germination in dormant and partially afterripened dormant caryopses of Avena fatua L. The rate of caryopsis germination was slower and final percentage germination lower in the highly dormant inbred line M73 at a given concentration of fusicoccin than in the dormant caryopses of line AN265. Gibberellic acid was more effective than fusicoccin in breaking dormancy in both lines. Promotion of germination of dormant caryopses by fusicoccin was inhibited by a 6-day pretreatment with (2-chloroethyl)trimethylammonium chloride.
The basal rate of proton efflux from embryos isolated from dormant and fully afterripened line AN265 caryopses was similar. Addition of fusicoccin increased the rate of proton efflux from the isolated embryos of dormant and afterripened caryopses by nearly 400%. Gibberellic acid had no effect on the rate of proton extrusion. The uptake of ⁸⁶Rb⁺ in dormant and afterripened A. fatua embryos was similar after a 2 h uptake period. The addition of fusicoccin to the medium doubled the uptake of ⁸⁶Rb⁴ by dormant and afterripened embryos. Gibberelleic acid had no effect on the uptake of ⁸⁶Rb⁺ by isolated embryos from either dormant or afterripened caryopses. The experimental results indicate that gibberellic acid is more versatile in its action than fusicoccin, and gibberellic acid may facilitate dormant A. fatua caryopsis germination by stimulating mechanisms other than the direct H⁺ efflux and K⁺ uptake at the membrane level.     

Seed germination studies with Heteropogon contortus

Germination characteristics of Heteropogon contortus were investigated in germination cabinets and in the glasshouse using soil trays. Under the former a dual inhibitory system appeared to operate. On the one hand naked caryopses from seeds which were less than 4 months of age were inhibited from germination by a process apparently analogous to epicotyl dormancy. This could readily be broken by applying 1% gibberellic acid and partly broken by repeated wetting and drying. On the other hand between 4 and 12 months, naked caryopses germinated more readily than intact seeds (caryopsis plus glumes, lemmas, etc.). However, the dormancy factor apparently associated with these structures could not be detected when seed was germinated in soil in the glasshouse and it appeared that they are a necessary protection for the caryopsis under field conditions. The optimal temperature for germination was between 30 and 35°C constant. Alternating diurnal temperatures of less than this appeared to cause reduced germination, and no high-temperature stimulation was observed. Germination was not directly promoted by the passage of a normal grass fire.     

Phytochrome-mediated germination control of maize caryopses

Maize caryopses sown in water germinate equally well either in darkness or under any light regime. However, when they are imbibed in mannitol solutions, continuous far-red light proves to be strongly inhibitory on the final germination as compared to darkness. Similar but less pronounced inhibition is also exhibited by continuous red or blue light. Intermittent far-red light can partially substitute for continuous far-red light in inhibiting maize caryopsis germination, and its effect is reversed to the intermittent red light level when red light is given immediately after each far-red illumination. These results are interpreted as a proof of existence and involvement of phytochrome in the germination control of maize caryopses, though its manifestation is realized only under osmotic stress.Abbreviations D darkness - FR far-red - R red - B blue - c-FR, c-R, c-B continuous FR, R, B, resp. - i-FR, i-R intermittent FR, R, resp.     

Distribution of Cadmium,Lead, Nickel,and Strontium in Imbibing Maize Caryopses

Seed germination is tolerant to heavy metals apparently because the seed coat is impermeable to metal ions. However, it is not clear whether the seed coat is a universal barrier for all metals. In addition, depending on their physical and chemical properties, a distribution of various metals may differ within an imbibing caryopsis, and therefore they produce dissimilar effects on seed germination. The toxic effects of Cd(NO₃)₂, Pb(NO₃)₂, Ni(NO₃)₂, and Sr(NO₃)₂ were estimated from the germination rates of maize (Zea mays L.) caryopses following two-day incubation with these salts. The distribution of heavy metals and Sr was studied by histochemical methods based on the formation of colored complexes with dithizone (Cd and Pb), dimethylglyoxyme (Ni), and sodium rhodizonate (Sr). Although the metals under study did not affect maize radicle protrusion, they inhibited seed germination in the following order: Cd > Ni ≈ Pb > Sr. Cd and Pb accumulated mainly in the seed coat cells, but Sr and Ni in the embryo cells and in the cells of endosperm (Sr) and scutellum (Ni). Although Cd was found only in the seed coat, it was the strongest inhibitor of seed germination. Apparently, due to high toxicity, Cd exerted its inhibitory effect at the concentrations too low for histochemical assay. In spite of easy translocation across the seed coat of imbibing caryopses, Sr did not considerably inhibit radicle protrusion and seed germination, apparently because of its low toxicity and predominant localization in the apoplast of embryo and endosperm cells.__________Translated from Fiziologiya Rastenii, Vol. 52, No. 4, 2005, pp. 635–640.Original Russian Text Copyright © 2005 by Seregin, Kozhevnikova.     

Water uptake and distribution in non-dormant and dormant wild oat (Avena fatua L.) caryopses

The influence of seed coat modification and light quality onwater uptake and distribution in caryopses of dormant and non-dormantlines of wild oat (Avena fatua L.) was determined using NMRmicroimaging. Non-dormant seeds absorbed water more rapidlythan dormant seeds during imbibition on distilled water. Thiseffect was detected first in the embryo-scutellar region (8h) and later in the proximal endosperm (12 h). Cutting the testaand pericarp close to the embryo or scarification with KOH promotedrapid embryo/scutellum hydration and germination. Cutting atthe middle part of the caryopsis did not enhance embryo hydrationnor did it greatly improve germination. The sensitivity of waterdistribution to the phytochrome germination effect was examined.Significant differences in imbibitional water uptake by embryos-scutellumtissue were detected by 18 h following red-light (germinationpromoter) compared with far-red (germination inhibitor) treatment.The results indicated that both the rate and the sequence ofembryo/scutellum hydration were important in initiating germinationin dormant seeds. A refinement of the model that describes waterimbibition in wild oat seeds during the early stages of germinationis discussed. Key words: Water uptake, water distribution, Avena fatua, seed coat modification, light quality, dormant and non-dormant seeds     

Factors influencing seed dormancy and germination in sand,and seedling survival under desiccation,of Psammochloa villosa (Poaceae), inhabiting the moving sand dunes of Ordos,China

Psammochloa villosa (Trin.) Bor. (Poaceae), is distributed primarily in moving sand dunes of the Ordos Plateau, China. Freshly harvested caryopses (seeds) are in non-deep physiological dormancy (non-deep PD). Germination is slow and low and only over a narrow temperature range. A treatment of four weeks cold stratification at 3 to 5 °C in darkness was required to break non-deep PD, allowing germination to reach high percentages at higher temperatures and without light requirement. Rate and percentages of germination were increased by scarifying the caryopsis coat and by artificial removal of different proportions of the endosperm. However, seedling dry weight and increases in root and shoot lengths, were significantly influenced by the proportion of the endosperm that remained on the caryopses. Higher percentages of seedling emergence were obtained from the shallowly buried caryopses, ranging from 0.5–2.0 cm, and the depth of the caryopses in the sand affected the time of germination. The deeper the caryopses were buried, the more that remained ungerminated and in enforced dormancy. The caryopses germinated when the upper sand layer was removed and buried caryopses were at a suitable sand depth for germination, or when the sand was aerated. In natural habitats, germinated seedlings may be wholly exposed to the air by sand erosion and thus be exposed to drought stress. However, young seedlings have the ability to tolerate desiccation and to recover after rehydration. Root length at the `point of no return' is 4 mm.     

Water uptake and splitting of wild oat caryopsis

Imbibition and germination experiments were conducted on the caryopses of wild oats (Avena fatua L.). The embryo envelopes, pericarp and aleurone layer, which completely cover the embryo-endosperm, do not form barriers against water uptake. The initial uptake of water is passive and the water moves across the pericarp with ease as it contains cracks; it is, however, transported across the aleurone layer through its cell walls into the endosperm and embryo of the caryopsis. The starchy endosperm enlarges due to water uptake causing the pericarp to rupture, thus exposing the aleuronelayer-covered seed. The aleurone layer is structurally heterogenous consistings of radially compressed irregular cells and cuboidal or radiallys tretched cells; the latter contains thicker walls. The former type is present along the abaxial side of the embryo and in the crease on the adaxial side of the caryopsis; the latter type covers the endosperm. The physical distention of the endosperm due to water uptake causes the rupture of the pericarp and the aleurone layer, and facilitates the emergence of the radicle and coleorhiza of the embryo during caryopsis germination.     

On dislodging caryopses of wild oats

The untwisting movement of the hygroscopic awn of wild oats (Avena fatua L.) causes the caryopses to rotate in anticlockwise direction and to dislodge from the spikelet. While untwisting, the awn generates more torque than is required to separate the caryopsis. The periodic shattering of caryopses caused by awn movement may be biologically significant in that the chances of having favorable moisture conditions for caryopses germination in the field at the time of dispersal are increased. Financial support by the Saskatchewan Research Council is gratefully acknowledged.     

beta-Amylases in Cereals : A Study of the Maize beta-Amylase System

β-Amylase of maize (Zea mays L.) caryopses was studied during development and germination by means of enzymic, electrophoretic, and immunochemical techniques. β-Amylase activity increased during caryopsis development to a maximum value at the beginning of the water content plateau (at this stage the enzyme was located primarily within the pericarp) and then decreased. Almost no β-amylase (activity or antigen) was found in either free or bound forms in the mature maize caryopsis. The activity increased again during seedling growth and reached much higher values. Both the aleurone layer (to a major extent) and the scutellum produced and secreted β-amylase during germination, the secretion being stimulated by Ca²⁺. No posttranslational modification of the enzyme was detected during germination. The molecular specific activity of the enzyme remained unchanged during the observed periods, indicating that the regulation of the activity is based essentially on protein turnover. The enzyme from developing and germinating caryopses was found to be identical in terms of antigenicity, isoelectric point, and molecular mass to the β-amylases extracted from the roots and the leaves of the maize seedling. The maize β-amylase resembles in all respects the ubiquitous β-amylase described for rye and wheat, whereas the major β-amylase of those cereals appears to be lacking in the maize caryopsis.     

The Effect of Temperature on Seed Set and Seed Development in Detached Spikelets of Perennial Ryegrass (Lolium perenne L.)

In perennial ryegrass seed yield is low. Pollination, fertilizationand early stages of seed development are critical periods forthe realization of the yield potential. As intact plants aredifficult to handle, we used detached spikelets to study effectsof temperature and genotype on seed set and seed development. Plants of four cultivars were grown in pots. Just prior to flowering,spikelets were detached from the ears and put on water. Afterpollination, various spikelets of each plant were placed atdifferent temperatures (14/14, 17/12 or 20/15°C). Sevendays after pollination the percentage seed set and the lengthof the developing seeds (caryopses) were assessed. Only floretsin the four basal positions were used. For each cultivar an analysis of variance for unbalanced datawas carried out. There were highly significant difference amonggenotypes for seed set and caryopsis length. There was no evidencefor genotypic variation for tolerance to low temperatures. Neithertemperature nor floret position had an effect on seed set. Thecaryopsis length, however, increased with increasing temperatureand basal florets produced longer caryopses than distal florets.The detached spikelets proved to be a useful experimental tool.Copyright1993, 1999 Academic Press Lolium perenne L., perennial ryegrass, seed yield, seed set, detached spikelets culture, temperature, floret position     

Morphology and Germination of Yellow and Brown Caryopses of Aegilops Geniculata Roth (Gramineae) Population from Italy

Abstract

The germination percentage of Aegilops geniculata Roth caryopses was studied on four Italian populations (Pisa, Manfredonia, Catania and Simbirizzi). Observation of the germination trend was carried out at constant temperatures of 10°, 20° and 30°C, from maturation up to 60 days after harvesting. Spike, spikelet and caryopsis phenotypic characteristics were also studied. The populations of Pisa, Manfredonia and Simbirizzi presented two different types of caryopses: yellow caryopses - larger, heavier and composing roughly 60% of total—and brown caryopses—smaller and lighter in weight. The Catania population formed an exception in that brown caryopses were virtually absent (5%). This Sicilian accession also showed the largest spikelet size.

Yellow caryopses germinated more rapidly than the dark ones, which were shown to be endowed with longer relative dormancy, above all at 10° and 30° C. Such a phenomenon allows the two types of caryopses to have a different germination ecology, thus providing A. geniculata with a broader choice of germination opportunities and consequently favouring spread and survival of the species.     

Dormancy Implications of Phosphorus Levels in Developing Caryopses of Wild Oats (Avena fatua L.)

The element phosphorus made up 0.5% of the dry weight of dehulled Avena fatua caryopses 7 days after anthesis (DAA), half of it inorganic (P_i). Caryopses detached and pierced 7 DAA germinated in vitro with a rapid drop in P_i levels. By 15–20 DAA caryopsis dry weight had increased three- to fourfold, but phosphorus made up less than 0.04% of the dry weight of this enlarged caryopsis. Caryopses at this stage germinated readily without piercing if incubated in vitro. A further decrease in P_i accompanied by a marked increase in phytate phosphorus began about 15 DAA and continued during later seed maturation. By 20 DAA, when embryos were relatively mature and endosperm cell division had ceased, a decrease in caryopsis water content (as a percentage of dry weight) began, and seed dormancy became apparent. As starch and phytate reserves accumulated, P_i and water levels of the caryopsis diminished. Higher levels of endogenous P_i coincided with the anabolic events of initial seed formation and, to a lesser extent, with anabolic events of seed germination. Decreasing P_i levels coincided with accumulation of nutrient reserves, lowering of water content, and the initiation of dormancy. The data suggest that (1) enzymes associated with the formation and development of the embryo may be activated by the high P_i levels present during initial seed differentiation; (2) embryo quiescence and dormancy are facilitated by the drop of P_i levels which accompanies the accumulation of starch and phytate reserves; and (3) the increase in P_i which accompanies seed afterripening aids in the termination of dormancy and the resumption of germination. Received August 15, 1996; accepted December 2, 1996     

Growth and development of embryo parts during the germination of caryopses of the wild oat (Avena fatua L.)

Wild oat (Avena fatua L.) caryopses were germinated on moist filter paper and under water in the presence and absence of hydrogen peroxide (H₂O₂). The sequential growth and development of embryo parts were studied. Germination, as indicated by radicle emergence, was least and slowest in caryopses submerged in deoxygenated water. The coleorhiza in such caryopses elongated much earlier than the root, in contrast to the other treatments where the coleorhiza and the root emerged at about the same time. In caryopses incubated on moist filter paper all embryo parts showed considerable growth. In H₂O₂ treated caryopses only the epicotyl showed substantial growth over the experimental period. In all treatments the first mitotic peaks were noticed at the same period. The occurrence of these early nuclear divisions may be due to release of 4 C nuclei from inhibition by the uptake of water during caryopsis imbibition. The mitosis continued in the radicle of the embryo in those caryopses germinating on moist filter paper, indicating occurrence of DNA synthesis. In the other two treatments, however, few divisions were detected. Here the early growth of the root, causing caryopsis germination, was due to cell elongation, especially in the proximal part of the root.     

Weedy adaptation in Setaria spp. IV. Changes in the germinative capacity of S. faberii (poaceae) embryos with development from anthesis to after abscission

Studies were conducted evaluating germinability states in giant foxtail (Setaria faberii) embryos, as well as surrounding tissues (hull, caryopsis), with germination assays. Further, seed age, fascicle arrangement, flowering patterns, and elongation in the inflorescence were evaluated. Both qualitatitive and quantitative morphological observations of the hull and the caryopsis were revealed by precisely determined fertilized spikelet age from anthesis until after seed abscission. Red coloration of the placental pad at ≈ 11 d after anthesis is probably a morphological indicator of physiological maturity. Germinability of giant foxtail embryos changed with development. Four qualitatively different types of embryo germination were observed during development of the seed: early disorganized callus growth at the basal, coleorhizal end of the embryo; germination of immature embryos with shortened and thickened axes; germination of the scutellum; and germination and growth of the coleoptile and coleorhiza in embryos aged 7 d after anthesis and older. Axis-specific embryo germinability was also observed. Inhibition of the embryo could be localized to the coleoptile, the coleorhiza, or both. These studies provide evidence for a complex model of germinability regulation based on the independent, asynchronous actions of the embryo, caryopsis, and hull compartments, as well as on their dependent, synchronous action. These studies provide evidence for a dynamic, developmental model of giant foxtail germinability regulation resulting in phenotypes with a wide range of germinability shed from an individual panicle. These diverse germinability phenotypes are found at all stages of development, but particularly when the seed is shed and the soil seed bank is replenished.     

The physiological basis of seed dormancy in Arena fatua V. Action of ethanol and other organic compounds

Ethanol (50-200 mM )induced germination in four genetically pure dormant lines of Avena fatua L. The sensitivity to this treatment was moderate immediately after harvest and increased steadily during six months of after-ripening. This sensitivity to ethanol was detectable much earlier during after-ripening than with two other germination promoters, NaN³, and NaNO³, Because ethanol can overcome dormancy in freshly harvested caryopses, the mode of action of ethanol in these caryopses apparently differs from that of the two other promoters, azide and nitrate. Nevertheless, it is clear that induction of germination by the three promoters is fully gibberellin-dependent since in each case this response can be blocked by the administration of 2-chlorocthyl trimethylammonium chloride, an inhibitor of gibberellin biosynthesis. Short-term incubation treatments with ethanol were relatively more effective than continuous treatments. These brief treatments were most effective when presented near the beginning of seed imbibition. Among other organic compounds tested only acetaldehyde significantly promoted germination in all lines tested. Propan-1-ol, butan-l-ol, chloral hydrate, procaine, methanol and chloroform were marginally effective on the least dormant lines, while ether, formaldehyde, acetone and ethyl acetate were ineffective. The mode of action of ethanol in overcoming dormancy in both freshly harvested and partly after-ripened caryopses is discussed and the possible role as a metabolic substrate or anaesthetic is indicated.     

ENDOGENOUS FLUCTUATIONS OF GERMINATION CAPACITY IN DACTYLIS GLOMERATA

Kummerow, Jochen. (U. Chile, Santiago.) Endogenous fluctuations of germination capacity in Dactylis glomerata. Amer. Jour. Bot. 50(9): 915–920. Illus. 1963.—Dactylis glomerata caryopses, ‘New Zealand’ strain, furnished by the Ministerio de Agricultura of Chile, were enclosed in test tubes, carefully sealed with a Bunsen flame, and stored under constant conditions. Their germination capacity was periodically determined. In 1958/1959, a 2-peak annual germination curve was obtained. The germination increase was related with a diminution of the water content of the caryopses. The same material was seeded in 8 different zones of the Chilean Central Valley between 28°34′ and 40°35′ S. lat. The same germination procedures were practiced with the caryopses harvested in these 8 parcels. In the 1960/1962 period, 1-peak germination curves were found. We did not find the same relation between water content and germination as found in 1958/ 1959. There exists a certain relation between the velocity of germination and caryopsis origin. The velocity of germination decreases from north to south.     

Effect of gibberellin on protein and non-structural carbohydrate in dormant Avena fatua caryopses

Application of gibberellic acid (GA₃) to dormant Avena fatua L. caryopses resulted in the termination of dormancy within 24 h as indicated by germination between 24 and 48 h. During the period of imbibition from 0 to 24 and 24 to 48 h changes occurred in protein and carbohydrate metabolism in GA-treated and untreated caryopses. Germination did not occur in untreated caryopses, therefore physiological changes in these caryopses were not associated with the termination of dormancy. GA-treatment increased the concentration of soluble and SDS-extractable protein in the endosperm tissue by 4 and 5%, respectively, over the 24 h untreated material; no changes were apparent when the protein profiles of GA-treated and untreated tissues were compared by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE) 0, 24 and 48 h after imbibition. The concentration of hexose and sucrose in the GA-treated endosperm tissue increased 189 and 151 μmol, respectively, over the untreated material at 24 h. Gibberellic acid had no effect on starch metabolism in the endosperm tissue in the first 24 h, the period associated with the termination of dormancy. The concentration of hexose increased by 57 μmol and starch decreased by 80 μmol in the GA-treated embryo tissue within 24 h. Our results demonstrate that exogenously applied GA influences sucrose and hexose metabolism in the endosperm tissue. The specific effect of GA on starch and hexose metabolism in the dormant A. fatua caryopsis embryo tissue may be associated with the termination of dormancy.