Seed Germination Biology of the Narrowly Endemic Species Lesquerella stonensis (Brassicaceae)

Abstract Lesquerella stonensis (Brassicaceae) is an obligate winter annual endemic to a small portion of Rutherford County in the Central Basin of Tennessee, where it grows in disturbed habitats. This species forms a persistent seed bank, and seeds remain viable in the soil for at least 6 years. Seeds are dormant at maturity in May and are dispersed as soon as they ripen. Some of the seeds produced in the current year, as well as some of those in the persistent seed bank, afterripen during late spring and summer; others do not afterripen and thus remain dormant. Seeds require actual or simulated spring/summer temperatures to come out of dormancy. Germination occurs in September and October. Fully afterripened seeds germinate over a wide range of thermoperiods (15/6–35/20°C) and to a much higher percentage in light (14 h photoperiod) than in darkness. The optimum daily thermoperiod for germination was 30/15°C. Nondormant seeds that do not germinate in autumn are induced back into dormancy (secondary dormancy) by low temperatures (e.g., 5°C) during winter, and those that are dormant do not afterripen; thus seeds cannot germinate in spring. These seed dormancy/ germination characteristics of L. stonensis do not differ from those reported for some geographically widespread, weedy species of winter annuals and thus do not help account for the narrow endemism of this species.     

Germination Ecophysiology of the Mesic Deciduous Forest Herb Polemonium reptans var. reptans (Polemoniaceae)

Abstract Seeds of Polemonium reptans var. reptans , a perennial herb of mesic deciduous forests in eastern North America, mature in late May-early June, and a high percentage of them are dormant. Seeds afterripened (came out of dormancy) during summer when kept in a nylon bag under leaves in a nonheated greenhouse or on wet soil in a 30/15°C incubator. The optimum temperature for germination of nondormant seeds was a simulated October (20/10°C) regime. In germination phenology studies in the nonheated greenhouse, 20–30% of the seeds that eventually germinated did so in October, and the remainder germinated the following February and March. Since low (5°C) winter temperatures promote some afterripening (ca. 50%) and do not cause nondormant seeds to re-enter dormancy, seeds that fail to germinate in autumn may germinate in spring. Thus, the taxon has very little potential to form a persistent seed bank. The large spatulate embryos and ability of seeds to afterripen at high temperatures means that seeds of P. reptans var. reptans have nondeep physiological dormancy, unlike many herbaceous woodland species, which have morphophysiological dormancy.     

DORMANCY AND GERMINATION IN SEEDS OF COMMON RAGWEED WITH REFERENCE TO BEAL'S BURIED SEED EXPERIMENT

Common ragweed (Ambrosia artemisiifolia L.) was one of 19 herbaceous weedy species used by Beal in his buried viable seed experiment started in 1879. No seeds germinated during the first 35 years of the experiment when germination tests were performed in late spring, summer or early autumn. Germination did occur in seeds buried for 40 years when seeds were exhumed and tested for germination in early spring. Data obtained in more recent research provide the probable explanation for these results. Seeds of common ragweed that do not germinate in spring enter secondary dormancy by mid to late spring and will not germinate until dormancy is broken the following late autumn and winter. Thus, during the first 35 years of the experiment seeds were dormant when tested for germination, whereas seeds buried for 40 years were nondormant. Seeds buried 50 years or longer did not germinate when tested in spring, probably because they had lost viability and/or seeds germinated during burial and seedlings died.     

Seed Germination Ecology of the Cold Desert Annual Isatis violascens (Brassicaceae): Two Levels of Physiological Dormancy and Role of the Pericarp

The occurrence of various species of Brassicaceae with indehiscent fruits in the cold deserts of NW China suggests that there are adaptive advantages of this trait. We hypothesized that the pericarp of the single-seeded silicles of Isatis violascens restricts embryo expansion and thus prevents germination for 1 or more years. Thus, our aim was to investigate the role of the pericarp in seed dormancy and germination of this species. The effects of afterripening, treatment with gibberellic acid (GA₃) and cold stratification on seed dormancy-break were tested using intact silicles and isolated seeds, and germination phenology was monitored in an experimental garden. The pericarp has a role in mechanically inhibiting germination of fresh seeds and promotes germination of nondormant seeds, but it does not facilitate formation of a persistent seed bank. Seeds in silicles in watered soil began to germinate earlier in autumn and germinated to higher percentages than isolated seeds. Sixty-two percent of seeds in the buried silicles germinated by the end of the first spring, and only 3% remained nongerminated and viable. Twenty to twenty-five percent of the seeds have nondeep physiological dormancy (PD) and 75–80% intermediate PD. Seeds with nondeep PD afterripen in summer and germinate inside the silicles in autumn if the soil is moist. Afterripening during summer significantly decreased the amount of cold stratification required to break intermediate PD. The presence of both nondeep and intermediate PD in the seed cohort may be a bet-hedging strategy.     

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.     

Seasonal cycle of seed dormancy controls the recruitment of Butia odorata (ARECACEAE) seedlings in savanna‐like palm tree formations in southern Brazil

Butia odorata (Barb. Rodr.) Noblick is a palm tree that grows in savanna‐like formations in subtropical regions of South America, and whose regeneration is threatened by agricultural management. Its diaspores are dormant after dispersal which takes place during the summer and early autumn. The aim of this study was to investigate seasonal and microhabitat effects on the germination and seedling recruitment of this palm species. Diaspores were sown in the field, in both open lands and forest patches. During 2 years, we measured seed germination, viability and moisture, seedling emergence and germination response to warm stratification of those seeds that failed to germinate in the field. Germination was concentrated during the summer, when soil temperatures were highest, whilst seedling emergence peaked in the autumn and early winter, when temperature and humidity conditions became less extreme. In open lands, there were two pulses of germination (first and second summer), whilst in forest patches, a single pulse (second summer) was detected. Although overall germination did not differ between microhabitats, the percentage of seedling emergence from seeds that remained buried until the end of the experiment was almost twice as large in the forest patches compared with open areas. The viability of seeds declined over time, particularly in open areas. Laboratory‐induced warm stratification was found to act on seed dormancy release in a cyclic way, being far more effective on seeds retrieved from the field in spring–summer months than in those retrieved in the winter. This cyclic pattern of dormancy in B. odorata seeds results in major seedling recruitment after the summer, under wetter and cooler conditions, thus reducing mortality risk. This process can be enhanced by the presence of surrounding vegetation, which both increases seedling emergence and/or prolongs seed viability.     

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.     

Fruit and seed heteromorphism in the cold desert annual ephemeral Diptychocarpus strictus (Brassicaceae) and possible adaptive significance

Background and Aims

Diptychocarpus strictus is an annual ephemeral in the cold desert of northwest China that produces heteromorphic fruits and seeds. The primary aims of this study were to characterize the morphology and anatomy of fruits and seeds of this species and compare the role of fruit and seed hetermorphism in dispersal and germination.

Methods

Shape, size, mass and dispersal of siliques and seeds and the thickness of the mucilage layer on seeds were measured, and the anatomy of siliques and seeds, the role of seed mucilage in water absorption/dehydration, germination and adherence of seeds to soil particles, the role of pericarp of lower siliques in seed dormancy and seed after-ripening and germination phenology were studied using standard procedures.

Key Results

Plants produce dehiscent upper siliques with a thin pericarp containing seeds with large wings and a thick mucilage layer and indehiscent lower siliques with a thick pericarp containing nearly wingless seeds with a thin mucilage layer. The dispersal ability of seeds from the upper siliques was much greater than that of intact lower siliques. Mucilage increased the amount of water absorbed by seeds and decreased the rate of dehydration. Seeds with a thick mucilage layer adhered to soil particles much better than those with a thin mucilage layer or those from which mucilage had been removed. Fresh seeds were physiologically dormant and after-ripened during summer. Non-dormant seeds germinated to high percentages in light and in darkness. Germination of seeds from upper siliques is delayed until spring primarily by drought in summer and autumn, whereas the thick, indehiscent pericarp prevents germination for >1 year of seeds retained in lower siliques.

Conclusions

The life cycle of D. strictus is morphologically and physiologically adapted to the cold desert environment in time and space via a combination of characters associated with fruit and seed heteromorphism.     

PHYSIOLOGICAL ECOLOGY OF GERMINATION OF VIOLA RAFINESQUII

Seeds of the winter annual Viola rafinesquii Greene exhibit true dormancy at the time of maturity and dispersal in mid to late spring. During the summer rest period the seeds pass from a state of true dormancy to one of relative dormancy and finally to what may be called a state of complete nondormancy. As the seeds enter relative dormancy they will germinate mostly at relatively low temperatures (10, 15, 15/6, and 20/10 C), but as after-ripening continues they gain the ability also to germinate at higher temperatures (20, 25, and 30/15 C). During June, July, and August seeds will not germinate at field temperatures even if kept continuously moist. But by September and October seeds may germinate to high percentages over a wide range of temperatures, including September and October field temperatures. This pattern of germination responses, involving breaking of true dormancy and widening of the temperature range for germination during relative dormancy, appears to be an adaptation of the species to a hot, dry season. Seeds of V. rafinesquii stored on continuously wet soil (field capacity) or on soil that was alternately wet and dried during the summer did not after-ripen at low temperatures (10, 15, 15/6, and 20/10 C) but did after-ripen fully at high temperatures (20, 25, 30/15, and 35/20 C). Thus, the high temperatures that V. rafinesquii “avoids” by passing the summer in the dormant seed stage actually are required to break seed dormancy and, therefore, are essential for completion of its life cycle.     

Annual dormancy cycles in buried seeds of shrub species: germination ecology of Sideritis serrata (Labiatae)

The germination ecology of Sideritis serrata was investigated in order to improve ex‐situ propagation techniques and management of their habitat. Specifically, we analysed: (i) influence of temperature, light conditions and seed age on germination patterns; (ii) phenology of germination; (iii) germinative response of buried seeds to seasonal temperature changes; (iv) temperature requirements for induction and breaking of secondary dormancy; (v) ability to form persistent soil seed banks; and (vi) seed bank dynamics. Freshly matured seeds showed conditional physiological dormancy, germinating at low and cool temperatures but not at high ones (28/14 and 32/18 °C). Germination ability increased with time of dry storage, suggesting the existence of non‐deep physiological dormancy. Under unheated shade‐house conditions, germination was concentrated in the first autumn. S. serrata seeds buried and exposed to natural seasonal temperature variations in the shade‐house, exhibited an annual conditional dormancy/non‐dormancy cycle, coming out of conditional dormancy in summer and re‐entering it in winter. Non‐dormant seeds were clearly induced into dormancy when stratified at 5 or 15/4 °C for 8 weeks. Dormant seeds, stratified at 28/14 or 32/18 °C for 16 weeks, became non‐dormant if they were subsequently incubated over a temperature range from 15/4 to 32/18 °C. S. serrata is able to form small persistent soil seed banks. The maximum seed life span in the soil was 4 years, decreasing with burial depth. This is the second report of an annual conditional dormancy/non‐dormancy cycle in seeds of shrub species.     

Temperature Response Pattern during Afterripening of Achenes of the Winter Annual Krigia oppositifolia (Asteraceae)

Abstract Freshly-matured achenes of Krigia oppositifolia Raf. were buried in soil at near-natural temperatures for 0–35 months and then exhumed and tested in light and darkness at (12/12 hr) daily thermoperiods of 15/6, 20/10, 25/15, 30/15 and 35/20°C. Achenes required light for germination and exhibited an annual dormancy/nondormancy cycle, being dormant in spring and nondormant in autumn. High summer temperatures (30/15, 35/20°C) fully promoted afterripening, whereas low temperatures (5, 15/6°C) prevented it. As buried seeds came out of dormancy in summer, they first germinated at medium temperatures (20/10, 25/15°C), but with additional afterripening the maximum and minimum temperatures for germination increased and decreased, respectively. Thus, during afterripening, achenes exhibit type 3 temperature responses, which otherwise are known only in two perennial Asteraceae and one perennial Liliaceae. The physiological responses of achenes of K. oppositifolia are unlike those of most winter annuals, which have type 1 responses—i.e., the maximum temperature for germination increases during afterripening. Also, they are unlike the majority of Asteraceae, which have type 2 responses—i.e., the minimum temperature for germination decreases during afterripening. Type 1 responses, typical of most winter annuals, have yet to be reported in the Asteraceae.     

Temperature controls seed germination and dormancy in the European woodland herbaceous perennial Erythronium dens-canis (Liliaceae)

We examined the germination ecology and the temperature requirements for germination of Erythronium dens-canis, under both outdoor and laboratory conditions. E. dens-canis is a spring flowering woodland geophyte widely distributed across Europe. Germination phenology, including embryo development and radicle and cotyledon emergence, were investigated in a natural population growing in Northern Italy. Immediately after harvest, seeds of E. dens-canis were either sown on agar in the laboratory under simulated seasonal temperatures or placed in nylon mesh sachets and buried in the wild. Embryos, undifferentiated at the time of seed dispersal, grew during summer and autumn conditions in the laboratory and in the wild, culminating in radicle emergence in winter when temperatures fell to ≈ 5 °C. Emergence of cotyledons did not occur immediately after radicle emergence, but was delayed until the end of winter. Laboratory experiments showed that temperature is the main factor controlling dormancy and germination, with seeds becoming non-dormant only when given warmth, followed by cold stratification. Unlike seeds of E. dens-canis that germinate in winter, in other Erythronium species radicle emergence occurs in autumn, while in some it is delayed until seeds are transferred from winter to spring conditions. Our results suggest that there is genetic and environmental control of the expression of seed dormancy amongst Erythronium species, which is related to local climate.     

Seed germination and life history syndromes in the California chaparral

Syndromes are life history responses that are correlated to environmental regimes and are shared by a group of species (Stebbins, 1974). In the California chaparral there are two syndromes contrasted by the timing of seedling recruitment relative to wildfires. One syndrome, here called the fire-recruiter or refractory seed syndrome, includes species (both resprouting and non-resprouting) which share the feature that the timing of seedling establishment is specialized to the first rainy season after fire. Included are woody, suffrutescent and annual life forms but no geophytes have this syndrome. These species are linked by the characteristic that their seeds have a dormancy which is readily broken by environmental stimuli such as intense heat shock or chemicals leached from charred wood. Such seeds are referred to as “refractory” and dormancy, in some cases, is due to seed coat impermeability (such seeds are commonly called hardseeded), but in other cases the mechanism is unknown. Seeds of some may require cold stratification and/or light in addition to fire related stimuli. In the absence of fire related cues, a portion or all of a species’ seed pool remains dormant. Most have locally dispersed seeds that persist in the soil seed bank until the site burns. Dispersal of propagules is largely during spring and summer which facilitates the avoidance of flowering and fruiting during the summer and fall drought. Within a life form (e.g., shrub, suffrutescent, etc.), the seeds of these species have less mass than those of species with non-refractory seeds and this possibly reflects the environmental favorableness of the postfire environment for seedling establishment. Regardless of when fire occurs, germination is normally delayed until late winter or early spring. In the absence of fire, or other disturbance, opportunities for population expansion are largely lacking for species with this syndrome. The other syndrome, here called the fire-resister or non-refractory seed syndrome, includes species that are resilient to frequent fires (mostly by vegetative resprouting), but require fire-free periods for recruiting new seedlings. Included are shrubs, subshrubs, suffrutescents, lianas, geophytes and annuals. All are linked by the characteristic that their seeds germinate in the absence of cues related to wildfires. In many cases no form of seed dormancy is present and the seeds germinate soon after dispersal; consequently these species do not accumulate a persistent seed bank. Germination and seedling establishment is independent of fire and thus opportunities for population expansion are also independent of fire. The demographic pattern of seedling recruitment varies with the life form. For shrubs, seedling recruitment may be restricted to sites free of fire for periods of a hundred years or more. Recruitment appears to require relatively mesic conditions and this may account for the patchy distribution of these species within the matrix of relatively arid sites. Finding such sites has selected for propagules specialized for wind or animal dispersal; the majority are bird dispersed. These shrub species all disperse fruits in fall and winter and this may have been selected to take advantage of migratory birds as well as to time dispersal to the winter rains typical of the mediterranean-climate. Germination typically occurs within several weeks of the first fall or winter rains. Maturation of flowers and fruits during the summer and fall drought may account for the distribution of these species on more mesic sites. Seed mass of these species is large and this may have been selected to provide an advantage to seedlings establishing under the canopy of this dense shrub community.     

Dormancy and Impotency of Cocklebur Seeds: IV. Effects of Gibberellic Acid, Benzyladenine, Thiourea, and Potassium Nitrate on the Growth of Embryonic Axis and Cotyledon Segments

Germination of nondormant but impotent small cocklebur seeds (Xanthium pennsylvanicum Wallr.) was promoted profoundly with thiourea or benzyladenine, and slightly with gibberellic acid. Gibberellic acid was ineffective in causing the germination of dormant cocklebur seeds, although thiourea and benzyladenine were effective. Experiments with excised seed pieces showed that the promotive effects of thiourea, benzyladenine, and gibberellic acid on cocklebur seed germination were associated with the enhancement of growth of seed parts; thiourea stimulated predominantly the axial growth, whereas benzyladenine stimulated predominantly the cotyledonary growth.     

Germination ecology and seed population dynamics of Digitalis purpurea

Summary The germination ecology and the dynamics of the generative reproduction in populations of Digitalis purpurea L. were investigated in the field as well as in experiments. Germination of fresh seeds in the dark on moist filter paper appeared to differ between populations. These differences were eliminated when a moist natural soil functioned as germination substrate. An interaction between the spectral composition of light and the germination substrate was present. Germination in gradients of light, temperature and soil moisture revealed some clear-cut results. Germination proved to be strongly dependent on the percentage of vegetation cover. During two years of burial in litter bags, the number of buried viable seeds did not decrease. From one generation of seeds produced in a natural population, 18% was introduced into the buried seed bank, 10% germinated in autumn and 24% was present as a enforced dormant surface seed bank in late autumn.The results are discussed in relation to secondary succession. can be derived from Milton (1936), Salisbury (1942) and Thompson and Grime (1979). Soil disturbance and germination seem to be correlated in D. purpurea (Grime 1979). The purpose of this study is to analyse the dormancy and germination behaviour of D. purpurea in relation to the relevant environmental factors in order to explain the mechanisms of entry into, and the escape of D. purpurea seeds from a seed bank. Furthermore, an attempt will be made to quantify seed rain as well as the fate of different germinating and non-germinating seed rain fractions in space and time per unit area, in different stages of succession.     

Parental overwintering history affects the responses of Thlaspi arvense to warming winters in the North

The overwintering conditions of northern plants are expected to change substantially due to global warming. For perennial plants, winter warming may increase the risk of frost damage if the plants start dehardening prematurely. On the other hand, evergreen plants may remain photosynthetically active and thereby benefit from milder winters. The positive and negative effects of mild winters on annual plants remain, however, largely unknown. We postulated that summer annuals may be susceptible to frost damage if the seeds germinate during a mild spell in winter. Winter annuals may utilize a warm period for photosynthesis. These questions were addressed in two consecutive experiments in which pot-grown individuals of Thlaspi arvense that overwintered in the field were exposed to an elevated temperature for 8 days in growth chambers in mid-winter. No premature germination was observed in summer annuals. However, in accordance with our hypothesis, winter annuals started photosynthesising very rapidly upon exposure to elevated temperature. The winter warming treatment affected neither the total number of seeds produced nor the mean seed weight. These seeds, possessing divergent parental overwintering histories, were used as starting material for the second experiment. Seeds originating from both summer and winter annual plants germinated both in the autumn and in the following spring. We observed a major parental effect associated with the winter warming treatment. The warm spell experienced by the mother plant (either as a winter annual rosette or as a summer annual seed) reduced the proportion of autumn germination in the next generation. Only 43% of the seeds of summer annuals possessing a parental warming history germinated before the winter, whereas the germination percentage of seeds with no previous winter warming history was 71%. In the case of seeds collected from winter annual plants, 4% of the seeds germinated in autumn if the mother plants experienced the warming treatment during the previous winter, whereas the corresponding value was 37% if the mother plants did not experience warming. Our results show that summer and winter annual individuals show diverse responses to warm spells in winter. Since the responses are not limited only to the generation that actually experiences the warm spell, but also appear in their offspring, long-term studies consisting of several generations are called for.     

ENVIRONMENTAL CONTROL OF SEED GERMINATION IN THLASPI ARVENSE (CRUCIFERAE)

The effect of environmental conditions during storage and imbibition on germination was investigated in field pennycress (Thlaspi arvense L.), a weed species that can behave as a winter or a summer annual. Freshly harvested seeds of an inbred line with a cold requirement for flowering exhibited primary dormancy that was rapidly lost following 1 month of afterripening in a dry state. Nondormant seeds were positively photoblastic. The light effect was mediated through phytochrome since germination was promoted by red light and inhibited by far red light. Seedling emergence was also inhibited by light filtered through a canopy of wheat leaves. Germination of field pennycress seeds was considerably more sensitive to moisture stress than two sympatric species, wild oat (Avena fatua L.) and wheat (Triticum aestivum L., cv. ERA). Seeds of the latter two species were chosen in order to compare the effect of water potential on germination in field pennycress with that in sympatric species. It was concluded that the major environmental factor limiting nondormant field pennycress seeds on the soil surface was water availability. Imbibition of fully afterripened seeds at low temperatures (6 C) induced a deep secondary dormancy. In contrast to primary dormancy, cold-induced dormancy was not alleviated by red light, alternating temperatures (21/5 C), or 2 months of dry storage at 6, 15, or 35 C. However, exogenous gibberellin A₃ or 24 weeks of dry storage resulted in germination in cold-induced dormant seeds. Secondary dormancy was not observed in fully afterripened seeds that were preincubated at 21 C for 1 or 2 days prior to the cold treatment. These results may explain the failure in field experiments to observe the cold-induced secondary dormancy that limits spring emergence in other winter annuals (J. Baskin, C. Baskin, Weed Res. 1979 19: 285–292).     

GERMINATION ECOLOGY OF PHACELIA DUBIA VAR. DUBIA IN TENNESSEE GLADES

The germination characteristics of a population of the winter annual Phacelia dubia (L.) Trel. var. dubia from the middle Tennessee cedar glades were investigated in an attempt to define the factor(s) regulating germination in nature. Factors considered were changes in physiological response of the seeds (after-ripening), temperature, age, light and darkness, and soil moisture. At seed dispersal (late May to early June), approximately 50 % of the seeds were non-dormant but, would germinate only at low temperatures (10–15 C). As the seeds aged from June to September, there was an increase in rate and total percent of germination at 10, 15, and 20 C, and the maximum temperature for germination increased to 25 C. Little or no germination occurred at the June, July, and August temperatures in 0- to 2-month-old seeds, even in seeds on soil that was kept continuously moist during this 3-month period. At the September, October, and November temperatures 3- to 5-month-old seeds germinated to high percentages. In all experiments seeds germinated better at a 14-hr photoperiod than in constant darkness. Inability of 0- to 2-month-old seeds to germinate at high summer temperatures allows P. dubia dubia to pass the dry summer in the seed stage, while increase in optimum and maximum temperatures for germination during the summer permits seeds to germinate in late summer and early fall when conditions are favorable for seedling survival and eventual maturation.     

Dormancy-breaking and germination requirements for seeds of Symphoricarpos orbiculatus (Caprifoliaceae)

Fruits (drupes) of Symphoricarpos orbiculatus ripen in autumn and are dispersed from autumn to spring. Seeds (true seed plus fibrous endocarp) are dormant at maturity, and they have a small, linear embryo that is underdeveloped. In contrast to previous reports, the endocarp and seed coat of S. orbiculatus are permeable to water; thus, seeds do not have physical dormancy. No fresh seeds germinated during 2 wk of incubation over a 15°/6°-35°/20°C range of thermoperiods in light (14-h photoperiod); gibberellic acid and warm or cold stratification alone did not overcome dormancy. One hundred percent of the seeds incubated in a simulated summer → autumn → winter → spring sequence of temperature regimes germinated, whereas none of those subjected to a winter → spring sequence did so. That is, cold stratification is effective in breaking dormancy only after seeds first are exposed to a period of warm temperatures. Likewise, embryos grew at cold temperatures only after seeds were exposed to warm temperatures. Thus, the seeds of S. orbiculatus have nondeep complex morphophysiological dormancy. As a result of dispersal phenology and dormancy-breaking requirements, in nature most seeds that germinate do so the second spring following maturity; a low to moderate percentage of the seeds may germinate the third spring. Seeds can germinate to high percentages under Quercus leaf litter and while buried in soil; they have little or no potential to form a long-lived soil seed bank.     

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.