MECHANISM OF SEED DORMANCY IN AMBROSIA ARTEMISIIFOLIA

Dormancy in Ambrosia artemisiifolia seeds was broken by 8 weeks of stratification. Germination of nondormant seeds was greater in light than in continuous darkness. Embryos of freshly harvested seeds were nondormant. Leaching and scarification did not stimulate germination of the dormant seeds. Exogenous gibberellin (GA₃) slightly increased germination of intact dormant seeds, and the effect was greatly increased by scarification. Germination was greater in the light in both tests. Exogenous indoleacetic acid did not stimulate germination of dormant seeds. Endogenous gibberellin and auxin content increased during stratification, and there was also a significant increase in GA during post-stratification at a favorable germination temperature. Inhibitors in the dormant seeds decreased during stratification and post-stratification. The high concentration of chlorogenic acid present in dormant seeds increased slightly during stratification. An unknown phenol very similar to chlorogenic acid in fluorescence and U.V. absorption significantly increased after 2 weeks of stratification. A significant decrease in the concentration of a second unidentified phenol occurred after 2 weeks of stratification. It is proposed that dormancy in Ambrosia artemisiifolia may be controlled by an inhibitor-promoter complex. The dormant seed is characterized by high inhibitor and low promoter levels. In the nondormant seed the balance was shifted to favor the promoter. Evidence suggests that the inhibitor involved may be abscisic acid and the promoters may be gibberellin and auxin. The content of auxin may be partially controlled by the concentration of phenols.     

EFFECT OF SEED DIMORPHISM ON THE DENSITY-DEPENDENT DYNAMICS OF EXPERIMENTAL POPULATIONS OF ATRIPLEX TRIANGULARIS (CHENOPODIACEAE)

The importance of seed size and density in determining individual plant performance and plant population dynamics in experimental populations of the halophyte Atriplex triangularis was studied. Two distinct seed morphs—large, light seeds and small, dark seeds—are produced by individual A. triangularis plants. Experimental populations consisting of seed size monocultures (large or small seeds) and seed size mixtures were established at three different densities, and the time of germination, plant size, plant survivorship, and plant fecundity were monitored. Marked variation in time of germination was observed among treatments and between seed sizes, but germination within any given treatment occurred over a five- to ten-day period. Large seeds produced larger plants than small seeds did, and this dichotomy was maintained over the course of the entire experiment. Germination date and seed size interacted such that larger plants grew from seeds which germinated earlier than those which germinated later, regardless of seed size. Germination date had a more pronounced effect than seed size did on plant mortality in high density populations. At high density, large seed monocultures experienced greater mortality than small seed monocultures did, but in seed size mixtures, the mortality was evenly distributed between plants from the two seed sizes. Regardless of density conditions and parentage, large and small seeds were produced in equal proportion by the plants. Total seed production, however, was dramatically affected by plant density, and to a lesser degree by germination date. Although seed size effects alone did not appear to affect directly final plant biomass and fecundity, effects of seed size early in ontogeny may have contributed to differences in fecundity.     

THE NINETY-YEAR PERIOD FOR DR. BEAL'S SEED VIABILITY EXPERIMENT

In 1879 Dr. W. J. Beal selected seeds of 23 different species of locally common plants, mixed 50 seeds of each species with moist sand in unstoppered one-pint bottles, and buried the bottles in a sandy knoll to be unearthed and the viability of the seeds tested periodically. The year 1970 marked the ninetieth year the seed had been buried, and the thirteenth bottle was recovered to test for seed viability. Of the three species which had germinated in the 1960 test (curly dock, Rumex crispus; evening primrose, Oenothera biennis; and moth mullein, Verbascum blattaria), only V. blattaria had viable seed with 20% germination. No other species germinated. All ten seedlings of V. blattaria were grown to maturity, and seeds were then harvested to study the possible deviations from normality and the requirements for seed germination. All seedlings emerging from the first progeny seed appeared normal. The most prominent requirement for germination was light, and this is a possible explanation of why the seeds had remained viable but dormant for so long a period. One-third of the freshly harvested seed germinated in darkness and, furthermore, redrying of dark-moistened seed in the absence of light induced additional germination. Germination of dark-moistened seed was not completely restored when the still moist seeds were subsequently exposed to light. However, when dark-moistened seeds were dried and then remoistened in the light, germination was about 95 %. About 5 % of the seed did not germinate under the conditions used. We find that 5 % of the population of V. blattaria seeds are dormant for unknown reasons, that 30 % will germinate if supplied only with moisture, and that 65 % are inhibited and require light and moisture simultaneously for germination. Supplying this 65 % of the population with moisture in darkness results in the development of a second type of inhibition which is no longer light reversible. It appears that the simultaneous requirement for light and moisture is an important factor in permitting V. blattaria seeds to remain dormant during prolonged burial.     

GERMINATION ECOLOGY OF SEDUM PULCHELLUM MICHX. (CRASSULACEAE)

Factors controlling the timing of seed germination were investigated in the small succulent winter annual Sedum pulchellum Michx. (Crassulaceae) in its natural habitat on unshaded limestone outcrops in northcentral Kentucky. At maturity in early July the dormant seeds are not dispersed but are retained in the fruits on the standing dead plants until September and October. Many, but not all, of the seeds afterripen in the fruits during summer, and at the time of dispersal some of them are dormant and some are nondormant. Germination and annual population establishment occur in September and October from seed reserves that have been in the soil for one or more years and from seeds produced in the current year. Germination of nondormant seeds may be prevented in autumn by lack of the appropriate combination of environmental factors including light, temperature and soil moisture in the seed's microsite. The effect of low winter temperatures on ungerminated seeds in the population is to induce nondormant seeds into secondary dormancy and to prevent afterripening of dormant seeds. Thus, in spring all the seeds in the population's seed reserve are dormant. During spring and summer some of these seeds afterripen, and they germinate in autumn when, and if, germination requirements are fulfilled.     

SEED DORMANCY AND GERMINATION IN MELAMPYRUM LINEARE

Immature seeds of Melampyrum lineare Desr. have very high germination percentages and dormancy is induced in a variable fraction of the seed crop during ripening. Correlated with this is the endogenous gibberellin-like activity which is found in considerable amounts in immature seeds, less in batches of ripe seeds, and is not detectable in batches containing only dormant seeds. For germination dormant seeds require activation followed by cold storage. In the laboratory activation is produced by allowing moist, dormant seeds to respire freely for several weeks at 20 C, or by treatment with exogenous GA₃. Dormancy appears to be most directly related to inability of the embryo to hydrolyze the thickened, mannan-containing endosperm cell walls. Embryos excised from dormant seed can be grown on agar enriched with whole macerated dormant seeds or with the ethanol-extractable materials from these (mostly sucrose and a glycoside). However, dormant seed material does not support growth when extracted to remove benzene- and ethanol-soluble materials.     

Dormancy in Cereal Seeds: II. THE NATURE OF THE GASEOUS EXCHANGE IN IMBIBED BARLEY AND RICE SEEDS

When barley seeds imbibe water, the O₂ uptake of non-dormantseeds is considerably less than that of dormant seeds for atleast the first 6 h, irrespective of the rate at which the seedshad previously lost dormancy. During the initial 6 h of imbibition, the CO₂ output of dormantbarley seeds is usually only slightly greater than and sometimesno different from that of nondormant seeds. The CO₂ output ofdormant seeds is reduced by about 66 percent by millimolar KCN,whereas that of non-dormant seeds is decreased by about 12–13per cent only. The CO₂ output of dormant barley in nitrogenis considerably less than the CO₂ output of non-dormant seedsunder the same conditions. Dormant rice seeds also show a higher initial O₂ uptake thannon-dormant seeds, though this is not generally as marked asin barley. Similarly, the initial CO₂ output of dormant seedsis distinctly greater than that of non-dormant seeds, but inmillimolar KCN it is depressed to a greater extent than in non-dormantseeds. In nitrogen, the CO₂ outputs of dormant and non-dormantseeds were found to be the same. Consequently, unlike barley,dormant rice seeds appear to be as capable of carrying out alcoholicfermentation under anaerobic conditions as nondormant seeds. In barley, increasing the O₂ tension from 21 per cent to 100per cent increased the oxygen uptake of dormant seeds more thanthat of non-dormant seeds (an increase of 53 per cent as against20–23 Per cent). In dormant seeds there was a concomitantincrease in CO₂ output (about 50 per cent), but the CO₂ outputof non-dormant seeds was hardly affected. High concentrations of CO₂ are inhibitory to the germinationof both dormant and non-dormant barley seeds. At a concentrationof 10 per cent, however, CO₂ is inhibitory only to dormant seeds,although at 2.5–5 per cent it is sometimes stimulatoryto the germination of dormant seeds. A 24–h treatmentwith appropriate concentrations of ethanol, lactic acid, oracetaldehyde is also stimulatory to the germination of dormantbarley seeds. Histochemical investigations in barley indicated the presenceof peroxidase, cytochrome oxidase, and -glycero-phosphate dehydrogenasein the embryo, aleurone layer, and in a layer associated withthe testa. A number of other redox enzymes were detected inthe embryo and aleurone layer only. No differences in distributionor intensity of activity were detected between dormant and nondormantseeds.     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

Evidence for physiological seed dormancy cycling in the woody shrub Asterolasia buxifolia and its ecological significance in fire‐prone systems

• Dormancy cycling is a key mechanism that contributes to the maintenance of long‐term persistent soil seed banks, but has not been recorded in long‐lived woody shrub species from fire‐prone environments. Such species rely on seed banks and dormancy break as important processes for post‐fire recruitment and recovery.
• We used germination experiments with smoke treatments on fresh seeds and those buried for 1 year (retrieved in spring) and 1.5 years (retrieved the following late autumn) to investigate whether Asterolasia buxifolia, a shrub from fire‐prone south‐eastern Australia with physiologically dormant seeds, exhibited dormancy cycling.
• All seeds had an obligation for winter seasonal temperatures and smoke to promote germination, even after ageing in the soil. A high proportion of germination was recorded from fresh seeds. but germination after the first retrieval was significantly lower, despite high seed viability. After the second retrieval, germination returned to the initial level. This indicates a pattern of annual dormancy cycling; one of the few observations, to our knowledge, for a perennial species. Additionally, A. buxifolia’s winter temperature and smoke requirements did not change over time, highlighting the potential for seeds to remain conditionally dormant (i.e. restricted to a narrow range of germination conditions) for long periods.
• For physiologically dormant species, such as A. buxifolia, we conclude that dormancy cycling is an important driver of successful regeneration, allowing seed bank persistence, sometimes for decades, during fire‐free periods unsuitable for successful recruitment, while ensuring that a large proportion of seeds are available for recruitment when a fire occurs.

     

The effect of extended exposure to hypersaline conditions on the germination of five inland halophyte species

In order to determine how salinity and exposure time affect seed viability and germination, seeds of five halophytes, Atriplex prostrata, Hordeum jubatum, Salicornia europaea, Spergularia marina, and Suaeda calceoliformis were exposed to 3.0, 5.0, and 10.0% NaCl solutions for 30, 60, 90, 365, and 730 d. Recovery experiments in distilled water indicated significantly different species responses to salinity over time. Percentage germination and rate of germination in H. jubatum were dramatically reduced following extended exposure and all seeds exposed to 10% NaCl for > 1 yr failed to germinate. Spergularia marina seeds were stimulated following short-term exposure to 3% NaCl; however, germination was delayed and overall germination was significantly reduced with exposure time in the two higher salinity levels. Percentage germination in A. prostrata decreased over time, but salinity level was not related to this reduction. Germination of S. europaea and S. calceoliformis, the most salt-tolerant species being tested, was stimulated by exposure to high salinity. Both species had a significant increase in percentage germination and in the germination rate when compared to seeds germinated in distilled water. Baseline germination data from seeds placed in 0, 1, 2, and 3% NaCl solutions indicated that S. europaea and S. calceoliformis were the only species to germinate in the 3% NaCl solution. Spergularia marina failed to germinate in the 2% NaCl treatment, and germination of A. prostrata and H. jubatum was significantly reduced at this salinity level. It is concluded that prolonged exposure to saline solutions can inhibit or stimulate germination in certain species, and the resulting germination and recovery responses are related to the duration and intensity of their exposure to salt in their natural habitats.     

Sensitivity to Seed Germination Schedule by Scatter‐Hoarding Pére David's Rock Squirrels During Mast and Non‐Mast Years

Mast seeding by oaks (Quercus) and other seed species has fundamental impacts on the behavior of individual scatter‐hoarding animals as well as their population and community dynamics. Scatter‐hoarding squirrels are highly sensitive to acorn germination schedule and stop germination by removing the embryo of non‐dormant acorns to prevent losses of energy and nutrients by rapid germination. However, we know little about how this behavior is affected by mast seeding. We investigated foraging decisions made by free‐ranging Pére David's rock squirrel (Sciurotamias davidianus) with three species that produce non‐dormant seeds and two species that produce dormant seeds (Fagaceae) during mast and non‐mast years in Central China from 2007–2010. Consistent with the food perishability hypothesis that squirrels make decisions to minimize the perishability of their caches, non‐dormant seeds were hoarded at a lower rate, but had a higher probability of embryo removal than dormant seeds. Compared with non‐mast years, a lower proportion of seeds were harvested during mast years, but a higher proportion of the harvested seeds were hoarded. In addition, the probability of embryo removal in the hoarded seeds was higher during mast years. Squirrels also dispersed seeds at shorter distances during mast years. Moreover, the interactions between mast seeding and seed germination schedule had significant effects on squirrel foraging decisions, including embryo removal. This study is the first to demonstrate that squirrels show greater sensitivity to seed germination potential when seeds are abundant.     

The effect of stratification temperatures on the level of dormancy in primary and secondary dormant seeds of two <Emphasis Type="Italic">Carex</Emphasis> species

The effect of temperature on the level of dormancy of primary and secondary dormant Carex pendula and Carex remota seeds was investigated. Primary dormant and secondary dormant seeds were stratified for 4 weeks at 5, 11, 13, and 15 °C, respectively, and tested for germination at 15/5 °C in light. To obtain secondary dormant seeds, primary dormant seeds were stratified at 5 °C and afterwards at 25 °C for 4 weeks. Germination tests were carried out in water and in 25 μmol KNO₃-solution to examine differences in sensitivity to nitrate between seeds relieved from primary and secondary dormancy. In both species, seeds with primary and with induced secondary dormancy showed no significant differences in germination. The two sedges showed significant differences in the effect of stratification temperatures between primary and secondary dormant seeds. Primary dormant seeds of C. pendula showed high germination (>80%) in nitrate-solution after stratification at all temperatures, while only temperatures of 5, 11, and 13 °C led to higher germination in nitrate-solution in secondary dormant seeds. Germination percentages of primary and of secondary dormant C. pendula seeds in water increased to a higher extent only after stratification at 5 and 11 °C; stratification of 11 °C was more effective in secondary than in primary dormant seeds. The only temperature that relieved primary dormancy in C. remota seeds was 5 °C where germination in water and nitrate-solution was >90%. Germination of secondary dormant seeds was increased by stratification at 11 °C independent of the test solution but higher germination after stratification at 13 °C occurred only in nitrate-solution. The results support the existence of physiological differences in the regulation of primary and secondary dormancy by temperature, and in the reaction of nitrate, at least in C. remota.     

Physiology of oil seeds: I. Regulation of dormancy in virginia-type peanut seeds

The germination and ethylene production by dormant Virginia-type peanut seeds were observed in relation to phytohormone treatments that could conceivably release the dormancy of these seeds. A comparison was made between the effects of these treatments on the less dormant apical seeds and the more dormant basal seeds. Indole-3-acetic acid did not stimulate ethylene production by, or germination of, the dormant seeds to any extent. Gibberellic acid at 5 × 10⁻⁴ M stimulated ethylene production by apical seeds to 17 millimicroliters per hour and germination to only 40% above the control. The more dormant basal seeds were affected even less by gibberellic acid than the seeds. Ethylene gas at 8 microliters per liter stimulated germination to 85% above the control for both apical and basal seeds. At this ethylene concentration the physiology of the more dormant basal seeds was altered, so that they behaved in a manner similar to the inherently less dormant apical seeds. 2-Chloroethylphosphonic acid at 10⁻³ and 5 × 10⁻⁴ M provided results similar to ethylene gas. Both apical and basal seeds germinated 100% at 48 hours. Among the phytohormones tested in this study, ethylene gas produced the greatest germination at low concentrations, and it appears must directly related to initiating the reactions required for converting the quiescent cells to an active state of growth.     

The Mechanism of Low Temperature Dormancy in Mature Seeds of Syringa Species

The mechanism of seed dormancy at low temperatures (15-9°C) was investigated in the seeds of Syringa josikaea, S. reflexa and S. vulgaris. Low temperature dormancy in Syringa species was mainly imposed by endosperm embedding the radicle. Different degrees of embryo dormancy may occur in S. reflexa seeds. In most cases the low temperature dormancy was broken completely by removing the endosperm around the radicle. The endosperm did not seem to contain significant quantities of germination inhibitors, and the results indicate that it prevents germination mainly due to its mechanical resistance. The mechanical resistance of endosperm did not change during chilling or during induction of dormancy by high temperature incubation. The strength of the endosperm decreased rapidly in non-dormant seeds before visible germination. Similar changes were not observed in dormant seeds. Generally, the strength of the endosperm was lower in the non- (or less) dormant species S. josikaea and S. vulgaris than in the more dormant S. reflexa seeds. The growth potential of the embryos, measured as their ability to germinate in osmotic solutions (mannitol or polyethylen glycol 4000), was increased by chilling and by GA₃-treatment. The growth potential of untreated S. josikaea and S. vulgaris embryos was generally higher than that of S. reflexa embryos. Acid ethyl acetate fractions of methanol extracts from embryos of all three species contained substances with GA³-like activity in the lettuce hypocotyl test. The activity was found at Rf 0.9–1.0 on paper chromatograms run in distilled water. No significant changes in the activity were detected during chilling or prior to visible germination.     

GERMINATION ECOPHYSIOLOGY OF ARENARIA GLABRA,A WINTER ANNUAL OF SANDSTONE AND GRANITE OUTCROPS OF SOUTHEASTERN UNITED STATES

The germination ecophysiology of Arenaria glabra Michx., a characteristic winter annual plant species of granite and sandstone outcrops of southeastern United States, was investigated. Seeds germinate in early autumn, plants overwinter in the rosette stage and then flower, set seeds, and die in late spring; seeds are dispersed soon after maturity. Eighty-five to 90% of freshly-matured seeds were innately dormant, and the other 10–15% germinated only at temperatures lower than those that occur in the habitat at the time of seed dispersal in June. During the summer after-ripening period, seeds stored dry under ambient laboratory conditions exhibited progressive increases in rates and total percentages of germination, a widening of the temperature range for germination, and a loss of the light requirement. At a 14-hr daily photoperiod, seeds kept on continuously moist soil germinated to 83% at simulated July and August temperatures during July and August, and the remainder germinated at September temperatures in September. On the other hand, seeds subjected to alternate wetting and drying during July and August germinated to only 9% during those 2 months, and the remainder germinated after the soil was kept continuously moist, beginning on 1 September, at simulated habitat temperatures during September and October. Thus, the timing of germination of A. glabra in the field is controlled by an interplay of the seeds' physiological state with the dynamics of temperature and soil moisture conditions.     

Effects of gut treatment on recovery and germinability of bovine and ovine ingested seeds of four woody species from the Sudanian savanna in West Africa

Passage rate through the digestive tracts of zebu cattle and sheep, and subsequent germination of egested seeds of four woody species from the Sudanian savanna, Acacia dudgeoni, Acacia seyal, Burkea africana and Prosopis africana, were studied. The result indicates large differences in passage rate among woody species, as well as between animals. The values ranged from 46% to 87% for seeds ingested by cattle while the lowest passage rate was 2.3% and the highest being 74% for seeds ingested by sheep. Among plant species, seeds of Prosopis africana had the highest passage rate through the digestive tract of both cattle and sheep. Seed passage through the gut showed a significant positive correlation with seed mass and thickness for cattle and sheep, respectively. The gut treatment and the retention time in the gut did not improve germination capacity and the speed of germination of dormant seeds. For non-dormant seeds of Acacia dudgeoni, the germination capacity was higher for seeds ingested by cattle than sheep. The speed of germination was also significantly higher for egested seeds than the control. It can be concluded that large herbivores could play an essential role in long distance dispersal of seeds. Gut treatment alone was not effective in breaking seed coat-imposed dormancy, although it enhanced the rate of germination of non-dormant seeds. To get a complete picture of the effect of frugivore on the release of seed dormancy, the combined effect of initial mastication and subsequent gut treatment needs to be investigated.     

Control of seed dormancy in Nicotiana plumbaginifolia: post-imbibition abscisic acid synthesis imposes dormancy maintenance

The physiological characteristics of seed dormancy in Nicotiana plumbaginifolia Viv. are described. The level of seed dormancy is defined by the delay in seed germination (i.e the time required prior to germination) under favourable environmental conditions. A wild-type line shows a clear primary dormancy, which is suppressed by afterripening, whereas an abscisic acid (ABA)-deficient mutant shows a non-dormant phenotype. We have investigated the role of ABA and gibberellic acid (GA₃) in the control of dormancy maintenance or breakage during imbibition in suitable conditions. It was found that fluridone, a carotenoid biosynthesis inhibitor, is almost as efficient as GA₃ in breaking dormancy. Dry dormant seeds contained more ABA than dry afterripened seeds and, during early imbibition, there was an accumulation of ABA in dormant seeds, but not in afterripened seeds. In addition, fluridone and exogenous GA₃ inhibited the accumulation of ABA in imbibed dormant seeds. This reveals an important role for ABA synthesis in dormancy maintenance in imbibed seeds. Received: 31 December 1998 / Accepted: 9 July 1999     

The role of invertebrates in seedling establishment in a heterocarpic plant,Atriplex sagittata

Seed heteromorphism is a marked character of many Chenopodioideae (Amaranthaceae). Seed morphs differ in dormancy, germination and seedling biology, but differences in their predation have not yet been studied. Atriplex sagittata produces small black dormant and large brown non‐dormant seeds. In this study, the timing of seed release and seedling establishment were ascertained, and their consumption by invertebrates (carabids, isopods and slugs) was studied. Seeds dispersed in the autumn passed the winter on the ground surface, protected from invertebrate predation by low temperatures. In the following vegetative season, ungerminated black seeds exposed to predation on ground surface were preferred by a large carabid species, Pseudoophonus rufipes. Some black and all brown seeds escaped predation by germinating in early spring. The seedlings were little endangered by carabids and isopods but were preferred by an invasive slug, Arion vulgaris, the feeding of which can exterminate seedlings at places in which slugs are abundant. Invertebrate predation is important factor of seed and seedling mortality of A. sagittata and seed heteromorphism modifies its intensity and timing.     

Maternal and paternal effects on the germination time of non‐dormant seeds of a monocarpic perennial species,Aster kantoensis (Compositae)

In plant species producing non‐dormant seeds, the germination time (from the start of imbibition to radicle emergence) is the main factor determining the timing of seedling emergence. We investigated maternal and paternal genetic effects on the germination time of non‐dormant seeds of a monocarpic perennial, Aster kantoensis Kitamura (Compositae). Three sets of reciprocal diallel crosses among five plants were conducted to produce genetic variation in seeds, and the germination time of the progeny of each parent was determined. The effects of the maternal parent and the interaction of maternal and paternal parents on the germination time of progeny were significant in all sets, and the effect of the paternal parent was significant in two of the three sets. This result means that the germination time of the progeny of a maternal or paternal parent can vary with the genotype of its mating partners. Because variation in the emergence time of seedlings contributes to avoiding seedling loss owing to unpredictable environmental changes, genetic variation in the germination time among the progeny of each parent mating with multiple partners could contribute to the establishment of the parent's seedlings in species producing non‐dormant seeds in the field.