Different requirements for physical dormancy release in two populations of <Emphasis Type="Italic">Sophora alopecuroides</Emphasis> relation to burial depth

Most species of Fabaceae produce seeds with physical dormancy which was broken by some environmental factors in the field, yet for the mechanism of physical dormancy release in the natural condition is still poorly understood. In present study, seeds of Sophora alopecuroides from two populations were placed on the soil surface or buried at depths of 2 and 7 cm in the field and were exhumed at time intervals to assess the changes of dormancy. Also, microenvironments were simulated in the laboratory to determine the possible components (temperature, temperature fluctuation, and moisture) involved in physical dormancy release. The laboratory work indicated that wet heat is an important dormancy release mechanism for both populations; and dry heat for one of them, but showed no effects on the other one. Consistent with that, burial experiment showed seeds of two populations had two contrasting-dormancy release patterns in the field. Most seeds of the dry heat sensitive population released dormancy but the other one remained dormant when seeds placed on the soil surface for 11 months (where seeds were exposed to dry heat but not wet heat). The difference of physical dormancy release pattern response to burial depth and simulated condition may attribute to seed coat color and hilum thickness which showed significantly different between two populations. This conclusion may improve understanding of physical dormancy release mechanism among populations and have a practical use for grassland management and weed control.     

Ethylene as a possible cue for seed germination of Schoenoplectus hallii (Cyperaceae), a rare summer annual of occasionally flooded sites

The purpose of our research was to determine why seeds of Schoenoplectus hallii germinate only in some wet years. Seeds mature in autumn, at which time they are dormant. Seeds come out of dormancy during winter, if buried in nonflooded, moist soil, but they remain dormant if buried in flooded soil. Nondormant seeds require flooding, light, and exposure to ethylene to germinate. One piece of apple in water (1/12 of an apple in 125 mL of water in a glass jar for a depth of 5 cm) or a 1-μmol/L solution of ethephon elicited very similar (high) germination percentages and vigor of seedlings. Apple, which was shown to produce ethylene in the air space of the jar, was used in a series of experiments to better understand germination. Seeds germinated to 72% if apple was removed from the water after 1 d of incubation, and they germinated to 97% if seeds were washed and placed in fresh water after 3 d of exposure to apple. No seeds germinated in control with no apple. Seeds incubated in apple leachate for 5 d and then transferred to filter paper moistened with distilled water germinated to 90%. Minimum depth of flooding in apple leachate (no soil in jars) for optimum germination was ≥3 cm. Buried seeds of S. hallii exhibited an annual conditional dormancy/nondormancy cycle. Regardless of the month in which seeds were exhumed, they germinated to 59-100% in light in water with apple at daily alternating temperature regimes of 25°/15°, 30°/15°, and 35°/20°C, but germination at 20°/10°C (and to some extent at 15°/6°C) tended to peak in autumn to spring. Thus, seeds can germinate throughout the summer if flooded (ethylene production) and exposed to light. An ethylene cue for germination serves as a "flood-detecting" mechanism and may serve as an indirect signal that water is available for completion of the life cycle and competing species are absent.     

Sensitivity of Polygonum aviculare seeds to light as affected by soil moisture conditions

BACKGROUND AND AIMS: It has been hypothesized that soil moisture conditions could affect the dormancy status of buried weed seeds, and, consequently, their sensitivity to light stimuli. In this study, an investigation is made of the effect of different soil moisture conditions during cold-induced dormancy loss on changes in the sensitivity of Polygonum aviculare seeds to light. METHODS: Seeds buried in pots were stored under different constant and fluctuating soil moisture environments at dormancy-releasing temperatures. Seeds were exhumed at regular intervals during storage and were exposed to different light treatments. Changes in the germination response of seeds to light treatments during storage under the different moisture environments were compared in order to determine the effect of soil moisture on the sensitivity to light of P. aviculare seeds. KEY RESULTS: Seed acquisition of low-fluence responses during dormancy release was not affected by either soil moisture fluctuations or different constant soil moisture contents. On the contrary, different soil moisture environments affected seed acquisition of very low fluence responses and the capacity of seeds to germinate in the dark. CONCLUSIONS: The results indicate that under field conditions, the sensitivity to light of buried weed seeds could be affected by the soil moisture environment experienced during the dormancy release season, and this could affect their emergence pattern.     

DORMANCY AND GERMINATION OF COMMON RAGWEED SEEDS IN THE FIELD

Common ragweed (Ambrosia artemisiifolia) seeds were stored under natural environmental conditions by placing them at three soil levels (surface, 5 cm, and 15 cm) in the field on November 1, 1972. Germination tests at 4-week intervals indicated that dormancy was broken by the end of January. Germination was initially greater at high temperatures, but this difference decreased with increasing time in the field. Secondary dormancy was evident in surface seeds by March 21 but not until April 18 at 5 cm and June 13 at 15 cm. Germination in the field was greatest at the surface but was observed at all soil levels by March 21. Seedling survival was 68% at the surface and 0% at 5 and 15 cm on June 13. Maximum and minimum soil temperatures were recorded at each soil level during the experiment and were correlated with the results. Greater germination and survival at the surface supports the evidence for ragweed's dependence on soil disturbance for germination, and the induction of secondary dormancy explains why ragweed does not constitute a dominant part of the vegetation when disturbance occurs after the soil warms to a critical point in the summer.     

The changing window of conditions that promotes germination of two fire ephemerals, Actinotus leucocephalus (Apiaceae) and Tersonia cyathiflora (Gyrostemonaceae)

BACKGROUND AND AIMS: Following a period of burial, more Actinotus leucocephalus (Apiaceae) and Tersonia cyathiflora (Gyrostemonaceae) seeds germinate in smoke water. The main aim of this study was to determine whether these fire-ephemeral seeds exhibit annual dormancy cycling during burial. This study also aimed to determine the effect of dormancy alleviation on the range of light and temperature conditions at which seeds germinate, and the possible factors driving changes in seed dormancy during burial. METHODS: Seeds were collected in summer, buried in soil in mesh bags in autumn and exhumed every 6 months for 24 months. Germination of exhumed and laboratory-stored (15 degrees C) seeds was assessed at 20 degrees C in water or smoke water. Germination response to light or dark conditions, incubation temperature (10, 15, 20, 25 and 30 degrees C), nitrate and gibberellic acid were also examined following burial or laboratory storage for 24 months. In the laboratory seeds were also stored at various temperatures (5, 15, 37 and 20/50 degrees C) for 1, 2 and 3 months followed by germination testing in water or smoke water. KEY RESULTS: The two species exhibited dormancy cycling during soil burial, producing low levels of germination in response to smoke water when exhumed in spring and high levels of germination in autumn. In autumn, seeds germinated in both light and dark and at a broader range of temperatures than did laboratory-stored seeds, and some Actinotus leucocephalus seeds also germinated in water alone. Dormancy release of Actinotus leucocephalus was slow during dry storage at 15 degrees C and more rapid at higher temperatures (37 and 20/50 degrees C); weekly wet/dry cycles further accelerated the rate of dormancy release. Cold stratification (5 degrees C) induced secondary dormancy. By contrast, no Tersonia cyathiflora seeds germinated following any of the laboratory storage treatments. CONCLUSIONS: Temperature and moisture influence dormancy cycling in Actinotus leucocephalus seeds. These factors alone did not simulate dormancy cycling of Tersonia cyathiflora seeds under the conditions tested.     

A combined physical and physiological dormancy controls seasonal seedling emergence of Geranium robertianum

Temperate forest herbs with seeds exhibiting both a physical and a physiological dormancy mechanism are rare, and knowledge on the factors regulating germination of these species is fragmentary. The biennial Geranium robertianum L. grows mainly in temperate woodlands, but can also be found in exposed habitats. Seedlings of G. robertianum are known to emerge from spring until autumn, but little is known about the environmental factors regulating germination. In this study, phenology of seedling emergence and of physical dormancy loss was examined for seeds buried at shaded or sunny exposed locations. The role of temperature in regulating dormancy and germination was analysed by incubating seeds in temperature sequences simulating temperatures that seeds experience in nature. The results indicate that most seeds of G. robertianum buried in sunny conditions germinate immediately after physical dormancy loss in summer. Seeds buried in shaded conditions also lose physical dormancy mainly during summer, but remain physiologically dormant and do not germinate until late winter or early spring. Besides physical dormancy, seeds of G. robertianum also initially have a high level of physiological dormancy, which is reduced during dry storage. Physiological dormancy is reduced through chilling in winter, thus enabling the seeds to germinate at low temperatures. We conclude that a complex combination of physical and physiological dormancy ensures that G. robertianum seeds germinate in summer at exposed sites and in early spring at shaded sites.     

沙埋和种子大小对固沙禾草沙鞭的种子萌发与幼苗出土的影响

该文研究了野外条件下不同深度的沙埋对沙鞭(Psammochloa villosa)种子萌发和幼苗出土的影响,以及温室条件下种子大小对不同深度沙埋后的种子萌发和幼苗出土的影响。结果表明,沙埋深度显著影响沙鞭的种子萌发率、幼苗出土率和种子休眠率。沙子表面的种子不能萌发。2 cm的浅层沙埋时的种子萌发率和幼苗出土率最高,1 cm 沙埋的种子萌发率和幼苗出土率次之。沙埋深度超过2 cm之后,沙鞭的种子萌发率和幼苗出土率与沙埋深度呈负相关。2 cm的种子休眠率最低。从2 ~12 cm,种子休眠率随着沙埋深度的增加而增加。在幼苗能够出土的深度(1~6 cm),幼苗首次出土所需的时间随着沙埋深度的增加而延长。种子大小对沙鞭的种子萌发率没有显著影响。但是在深层沙埋(6 cm)时,与小种子相比,大种子产生的幼苗的出土率较高。从2~6 cm,大种子形成的幼苗的茎长度都较长。     

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.     

The viability and germination characteristics of exhumed Solanum mauritianum seeds buried for different periods of time

The site, depth and duration of burial significantly influenced the viability and state of dormancy of Solanum mauritianum seeds. Burial at a depth of 15 cm was most effective in reducing the level of conditional dormancy. Secondary dormancy was not induced at any of the environmental (burial) sites when seeds were maintained at 15 cm, where light and temperature fluctuations were minimal. When buried at 4 cm or maintained on the soil surface secondary dormancy was induced, particularly at the inland sites where environmental conditions such as temperature and moisture were more extreme. Conditional dormancy could generally be overcome by incubating seeds at 15/30 °C in the light, even after prolonged burial at unfavourable germination conditions. Gibberellic acid (500 mg l^–1) was very effective in breaking secondary dormancy of seeds induced by storage under unfavourable conditions after burial. These results have important implications for the control of this week in commercial forests.     

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

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

     

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.     

Attributes of the seed bank after a century of primary succession on islands in Lake Hjälmaren, Sweden

1 A large number of islands was created when the water table of Lake Hjälmaren, south central Sweden, was lowered between 1882 and 1886. We have complete lists of vascular plant species for 40 of these islands from 1886, 1892, 1903–04, 1927–28 and 1984–85.
2 We have investigated the seed bank on nine of these islands and compared species composition at different soil depths with the species lists from the islands in 1886–1985, and with the present vegetation in the area of seed bank sampling. We have also investigated the distribution in the soil of seeds from species with different ecological attributes, including seed longevity, successional status, seed weight, seed form and species longevity.
3 Seeds in soil samples were allowed to germinate over the course of two summers with an intermediate cold storage. We found 1944 seeds representing 65 taxa. The mean seed density was 84 seeds dm ^–2 .
4 The similarity between the surface soil (0–3 cm) seed bank and the vegetation at the different vegetation analyses increased from 1886 to 1993. The similarity between the present vegetation and the seed bank decreased with increasing soil depth, and the soil at 12–15 cm had no species in common with the present vegetation. Several species now absent from the vegetation were found in the seed bank.
5 Deeply buried seeds came from early successional, annual species with long-term persistent and low-weight seeds, as expected from seed bank theories, but were slightly elongated, which was in contrast to theories. Spherical seeds were associated with the surface soil, as were short-lived and high-weight seeds from late successional, perennial species.     

Soil temperatures after the passage of a fire: Do they influence the germination of buried seeds?

Abstract Soil temperatures down to a depth of 5 cm were measured in the days following one fire in summer, one fire in winter and in unburnt vegetation during summer. Soil temperatures did not rise above 40°C after the winter fire or in unburnt vegetation during summer. Consequently, no impact on seed dormancy in the soil seedbank was expected. After a summer fire, soil temperatures above 40°C were found up to 4.5 cm in depth, while temperatures above 60°C were found only in the top 0.5 cm of soil. These temperatures are sufficient to break seed dormancy in some legume species in the seedbank. Hence, the season of burn may influence the number of seeds in the soil that have their dormancy broken and subsequent germination levels.     

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.     

Eastern gamagrass (Tripsacum dactyloides) root penetration into and chemical properties of claypan soils

Claypans restrict rooting depth and availability of moisture and nutrients to plants during periods of drought. Eastern gamagrass (Tripsacum dactyloides var. dactyloides [L.] L.) often remains green during summer droughts, while other plants turn brown. Questions arose whether eastern gamagrass roots had or could penetrate claypans to obtain needed moisture. Pits were dug (2 m deep) under eastern gamagrass plants that had been growing 50+ and 5+ years at two sites in Missouri. Clay contents were 30 to 50% in soil layers below 30 cm, and moisture was not limiting in these deep soil layers. Soil pH_Ca in the lower soil layers, except at 180 cm, was below 5.0, and in some cases near 4.0. Extractable Al was especially high in the 90 and 120 cm deep soil layers where pH was low. Extractable Ca, Mg, and K increased with soil depth. The eastern gamagrass roots effectively penetrated claypan soils. Root lengths and root weights were extensive to 180 cm depth, and decreased from the surface with soil depth. Roots of eastern gamagrass were aerenchymous (having cellular compartments which allow air movement) at all depths, were mycorrhizal to at least 150 cm depth, and had relatively high tolerance to acidic Al toxic Tatum subsoil (Typic Hapludult) and toxic levels of Al in nutrient solution. The eastern gamagrass roots also provided root channels through claypans, which could enable new eastern gamagrass or other plant roots to grow into deeper soil layers.     

埋藏条件下3种干旱荒漠植物的种子休眠释放和土壤种子库

对种子休眠的自然释放及其作用因素的研究, 是了解种子休眠生态学、种群适应机制的重要途径。以内蒙古阿拉善干旱荒漠区的3种主要植物牛枝子(Lespedeza potaninii)、唐古特白刺(Nitraria tangutorum)和骆驼蒿(Peganum nigellastrum)为材料, 研究了种子在野外埋藏18个月期间和4个埋深条件下的休眠释放特性和土壤种子库。3种植物种子在野外埋藏时(采收后5 ℃冷藏6个月)的休眠率分别为98%、95%和3%。结果显示, 埋藏过程中, 3种植物种子的休眠释放表现出不同的变化特性。对牛枝子而言, 置于地表(0 cm)的种子比埋藏于土中的种子的休眠释放快, 埋藏期末, 埋深0、2、5和10 cm的种子的休眠率分别为64%、87%、86%和82%。唐古特白刺种子埋藏6个月后, 各埋深的休眠已完全释放, 释放速率随埋深增加而加快。骆驼蒿种子具有典型的季节性休眠循环特性, 休眠率各年度最高点出现在10月份, 释放速率随埋深增加呈减慢趋势。埋藏期末不同埋深条件下, 牛枝子、唐古特白刺和骆驼蒿种子的平均田间萌发率分别为11%、12%和8%; 平均室内萌发率分别为3%、74%和42%; 而平均死种子率分别为3%、15%和10%。根据Thompson和Grime (1979)的土壤种子库分类体系, 供试的3种植物都属于持久土壤种子库类型。     

The responses of carrot fly larvae, Psila rosae, to components of their physical enrivonment

Abstract. 1. The responses of third instar Psila rosae (F.) larvae to light, temperature, humidity and soil moisture were investigated in the laboratory.
2. Larvae were photonegative and preferred a temperature of about 15°C. Temperatures between 30 and 40°C adversely affected movement and over 40° C were lethal.
3. In choice chambers, larvae preferred humidities of 70–100% r.h. and larvae in sand avoided dry conditions (2.5% field capacity). The latter response became more marked as larvae approached the pre-pupal stage when moistures of 40% field capacity and lower were avoided.
4. Most larvae were found at a depth of 8 cm in sand of uniform moisture content and temperature, but variation in moisture content could alter this preference.
5. In August, most larval damage in the field occurred near the tip of the carrot tap root but was more evenly distributed over the roots in November. It is uncertain whether this was due to soil near the surface being drier in August or whether it was caused by behavioural differences between the two generations of carrot fly larvae.
6. During the summer of 1975, low soil moisture levels resulted in the total absence of larval mines on the carrot roots even though pupae were found at depths of 20–30 cm in the soil. Temperature had no effect on the distribution of mines on carrot roots except at the top 2 cm of the soil profile.     

GERMINATION ECOPHYSIOLOGY OF HYDROPHYLLUM APPENDICULATUM,A MESIC FOREST BIENNIAL

In north central Kentucky, seeds of the mesic forest biennial Hydrophyllum appendiculatum Michx., are innately dormant at maturity in June. Under natural and simulated seasonal temperature changes, dormancy break occurred in two stages. Root dormancy was broken by high summer temperatures, and shoot dormancy was broken by low winter temperatures. Consequently, roots emerged from seeds during autumn, and cotyledons emerged the following spring. A 90-day warm (30/15 C) stratification treatment broke root dormancy, but the roots emerged only after transfer to lower temperatures. After the warm stratification treatment, roots emerged from 93, 73, 6 and 9% of the seeds incubated at 5, 15/6, 20/10 and 30/15 C (12/12 hr), respectively. Zero, 28, 56 and 84 days of cold (5 C) stratification of seeds with emerged roots resulted in 9, 21, 49 and 82% cotyledon emergence, respectively, at 20/10 C. Thus, H. appendiculatum exhibits a type of morpho-physiological dormancy known as epicotyl dormancy. Although many seeds germinate the first year, others remain dormant and germinate in successive years until the fourth season after ripening.     

Field fate of Amaranthus patulus seeds subjected to leaf-canopy inhibition of germination

Summary The germination of seeds of Amaranthus patulus Bertol., is known to be sensitive to leaf-transmitted light. Seeds were enclosed in transparent polyester-mesh envelopes and placed horizontally in 10-cm deep soil or on the soil surface, beneath a closed vegetation cover in the field. Changes in the numbers of firm intact seeds and of germinable seeds were traced for up to 3 years by periodical retrievals and germination tests. Rapid loss of germinable seeds, mainly due to germination, was observed in the buried seed population, in which only 20% of seeds maintained their germinability after 1 year, and a negligible number after 3 years. In contrast, the seeds placed on the soil surface maintained germinability relatively well: over 80% of seeds remained germinable after 1 year and a low percentage still preserved their germinability after 3 years. Assuming exponential decay in germinability, the decay rates on and in the soil were calculated from the data of the 1-year experiment to be 0.21 and 0.84 year^-1 respectively. The fate of seeds that were exposed to canopy light on the soil for a month and then buried was shown to be almost the same as that of the seeds which had been continuously in 10-cm deep soil. Correspondingly, the possibility of the induction of secondary (induced) dormancy by exposure to canopy light was excluded in a laboratory experiment, in which it was found that the imbibed seeds suffering leaf-canopy inhibition of germination exuded some diffusible germination inhibitor responsible for apparent dormancy. Estimation of numbers of A. patulus in the seed bank of an early successional field showed that 3,500 seeds/m² remained in the soil to the depth of 10 cm after 3 years' exclusion of the species following the production of 700,000 seeds/m², by a population explosively established after experimental induction of secondary succession.     

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.