Sequential temperature control of multi-phasic dormancy release and germination of Paeonia corsica seeds

Aims The physiological responses during dormancy removal and multi-phasic germination were investigated in seeds of Paeonia corsica (Paeoniaceae).Methods Seeds of P. corsica were incubated in the light at a range of temperatures (10–25 and 25/10°C), without any pre-treatment, after W (3 months at 25°C), C (3 months at 5°C) and W + C (3 months at 25°C followed by 3 months at 5°C) stratification, and a GA 3 treatment (250 mg·l^-1 in the germination substrate). Embryo growth, time from testa to endosperm rupture and radicle emergence were assessed as separate phases. Epicotyl–plumule emergence was evaluated incubating the germinated seeds at 15°C for 2 weeks, at 5 and 25°C for 2 months on agar water before transplanting to the soil substrate at 10, 15 and 20°C and at 15°C for 2 months on the surface agar water with GA 3 .Important findings Embryo growth, testa rupture, endosperm rupture (radicle emergence) and growth of the epicotyl were identified as four sequential steps in seeds of P. corsica. Gibberellic acid alone and warm stratification followed by 15°C promoted embryo growth and subsequent seed germination. Cold stratification induced secondary dormancy, even when applied after warm stratification. After radicle emergence, epicotyl–plumule emergence was delayed for ca. 3 months. Mean time of epicotyl–plumule emergence was positively affected by cold stratification (2 months at 5°C) and GA 3. P. corsica seeds exhibited differential temperature sensitivity for the four sequential steps in the removal of dormancy and germination processes that resulted in the precise and optimal timing of seedling emergence.     

冷层积和室温干燥贮藏对河西走廊8种荒漠植物种子萌发的影响

不同贮藏和处理条件对不同植物的种子萌发有不同的影响。该文以河西走廊干旱半干旱区8种荒漠植物为研究对象, 探讨了种子经历不同冷层积(4 ℃、-5 ℃、-26 - 10 ℃)和室温干燥贮藏后的萌发响应。研究结果表明: 1)冷层积可使种子萌发率提高、保持不变或降低, 冷层积的有效温度下界可降至-5 ℃或更低。4 ℃和-5 ℃的冷层积使多裂骆驼蓬(Peganum multisectum)和驼蹄瓣(Zygophyllum fabago)种子的萌发率升高、萌发速度加快, 冬季过低的气温以及较大的温度变幅(-26 - 10 ℃)使部分种子萌发率升高。3种冷层积和室温干燥贮藏使黑果枸杞(Lycium ruthenicum)种子萌发率达到90%-100%。唐古特白刺(Nitraria tangutorum)、甘草(Glycyrrhiza uralensis)、苦马豆(Sphaerophysa salsula)种子经过3种冷层积和室温干燥贮藏后萌发率变化较小。中亚紫菀木(Asterothamnus centrali-asiaticus)种子对各种贮藏条件的响应不明显, 部分种子活性丢失。刺沙蓬(Salsola ruthenica)种子扩散时有较高的萌发率(84%), 经-5 ℃和-26 - 10 ℃冷贮藏后, 种子仍具有较高的萌发率, 经4 ℃冷贮藏后几乎不萌发, 大部分种子活性丢失。2)不同物种的种子经过不同方式的贮藏后, 萌发对温度的响应不同。经冷层积后的多裂骆驼蓬种子萌发响应于恒温, 驼蹄瓣和刺沙蓬种子萌发更加响应于变温条件; 多数植物种子在变温培养下萌发速度慢于恒温下。     

Interchangeable effects of gibberellic acid and temperature on embryo growth, seed germination and epicotyl emergence in Ribes multiflorum ssp. sandalioticum (Grossulariaceae)

Morphophysiological dormancy was investigated in seeds of Ribes multiflorum Kit ex Roem et Schult. ssp. sandalioticum Arrigoni, a rare mountain species endemic to Sardinia (Italy). There were no differences in imbibition rates between intact and scarified seeds, suggesting a lack of physical dormancy, while methylene blue solution (0.5%) highlighted a preferential pathway for solution entrance through the raphe. Embryos were small at seed dispersal, with an initial embryo:seed ratio (E:S) of ca. 0.2 (embryo length, ca. 0.5 mm), whereas the critical E:S ratio for germination was three times longer (ca. 0.6). Gibberellic acid (GA(3), 250 mg · l(-1)) and warm stratification (25 °C for 3 months) followed by low temperature (<15 °C) enhanced embryo growth rate (maximum of ca. 0.04 mm · day(-1) at 10 °C) and subsequent seed germination (radicle emergence; ca. 80% at 10 °C). Low germination occurred at warmer temperatures, and cold stratification (5 °C for 3 months) induced secondary dormancy. After radicle emergence, epicotyl emergence was delayed for ca. 2 months for seeds from three different populations. Mean time of epicotyl emergence was affected by GA(3) . Seeds of this species showed non-deep simple (root) - non-deep simple (epicotyl) morphophysiological dormancy, highlighting a high synchronisation with Mediterranean seasonality in all the investigated populations.     

Temperature requirements differ for the two stages of seed dormancy break in Aegopodium podagraria (Apiaceae), a species with deep complex morphophysiological dormancy

Only a few studies have considered the possibility that low temperature requirements may vary among stages of dormancy break in seeds with morphophysiological dormancy (MPD). We show that this lack of consideration in previous studies on seed dormancy and germination of Aegopodium podagraria might explain the low germination percentages and/or the relatively long periods of incubation needed for germination. Under natural temperatures, embryos began to grow in September and were fully elongated by late December; most growth occurred when the average daily mean temperature was about 10°C. Radicles emerged under snow in late winter, and cotyledons emerged after snowmelt in early spring. In laboratory experiments, 100% of the embryos grew to full length at both 0 and 5°C, whereas 0°C was much more effective than 5°C in overcoming the physiological dormancy in seeds after embryos were fully elongated. Following radicle emergence, cotyledons emerged readily in a wide range of temperatures ≥5°C. GA(3) did not substitute for the low temperature requirement for dormancy break. Seed dormancy in A. podagraria fits Nikolaeva's formula for deep complex MPD, i.e., C(3)B-C(3). Better germination of seeds pretreated at 0° than at 5°C has practical implications for cultivating this species.     

Seed germination ecology of the threatened endemic Iberian <Emphasis Type="Italic">Delphinium fissum</Emphasis> subsp. <Emphasis Type="Italic">sordidum</Emphasis> (Ranunculaceae)

Seeds of Delphinium fissum subsp. sordidum are physiologically dormant at maturity, with underdeveloped embryos; thus they have morphophysiological dormancy (MPD). The aims of this study were to determine the requirements for embryo growth, dormancy break and germination, to characterise the type of seed dormancy and to evaluate the effects of light, seed age, pollination mechanism, and inter-annual and inter-population variability on germinative ability. After 3 months of incubation at 5°C (cold stratification) in darkness conditions, the mean embryo length increased from 5.6 to 2.07 mm, with 76% of seeds germinating. Conversely, embryos of seeds incubated during 3 months at 20/7 or 28/14°C hardly grew and no germination was recorded. Since cold stratification was the only requirement for the loss of MPD, and both dry storage in laboratory conditions and warm stratification prior to cold stratification shortened the cold stratification period required for germination, it could be concluded that D. fissum subsp. sordidum seeds have intermediate complex MPD. Cold stratification and incubation in darkness conditions promoted higher germination percentages than those in light. In addition, germinative ability increased with seed age up to 8 months (reaching 96% at 5°C in darkness), showed a pronounced inter-annual and inter-population variability, as well as a significant decrease in seeds coming from pollination by geitonogamy. High temperatures (25/10 or 28/14°C) induced seeds to secondary dormancy, so seedling emergence in the greenhouse was restricted to February–March. The requirements for dormancy break and germination reflect an adaptation to trigger germination in late winter. This study is the first one to document a gradual increase in germination percentage with seed age for plant species with intermediate complex MPD.     

Thermal thresholds as predictors of seed dormancy release and germination timing: altitude-related risks from climate warming for the wild grapevine Vitis vinifera subsp. sylvestris

Background and Aims

The importance of thermal thresholds for predicting seed dormancy release and germination timing under the present climate conditions and simulated climate change scenarios was investigated. In particular, Vitis vinifera subsp. sylvestris was investigated in four Sardinian populations over the full altitudinal range of the species (from approx. 100 to 800 m a.s.l).

Methods

Dried and fresh seeds from each population were incubated in the light at a range of temperatures (10–25 and 25/10 °C), without any pre-treatment and after a warm (3 months at 25 °C) or a cold (3 months at 5 °C) stratification. A thermal time approach was then applied to the germination results for dried seeds and the seed responses were modelled according to the present climate conditions and two simulated scenarios of the Intergovernmental Panel on Climate Change (IPCC): B1 (+1·8 °C) and A2 (+3·4 °C).

Key Results

Cold stratification released physiological dormancy, while very few seeds germinated without treatments or after warm stratification. Fresh, cold-stratified seeds germinated significantly better (>80 %) at temperatures ≥20 °C than at lower temperatures. A base temperature for germination (T_b) of 9·0–11·3 °C and a thermal time requirement for 50 % of germination (θ₅₀) ranging from 33·6 °Cd to 68·6 °Cd were identified for non-dormant cold-stratified seeds, depending on the populations. This complex combination of thermal requirements for dormancy release and germination allowed prediction of field emergence from March to May under the present climatic conditions for the investigated populations.

Conclusions

The thermal thresholds for seed germination identified in this study (T_b and θ₅₀) explained the differences in seed germination detected among populations. Under the two simulated IPCC scenarios, an altitude-related risk from climate warming is identified, with lowland populations being more threatened due to a compromised seed dormancy release and a narrowed seed germination window.     

Role of warm stratification in promoting germination of seeds of Empetrum hermaphroditum (Empetraceae), a circumboreal species with a stony endocarp

The broad objective of this research was to define the role of warm (≥15°C) stratification in breaking dormancy in seeds with stony endocarps that require warm-plus-cold (～0°-10°C) stratification for germination. This question was addressed using seeds (true seed + endocarp, hereafter called seeds) of Empetrum hermaphroditum. Only 2-5% of freshly matured seeds collected in September and October at five sites in Sweden germinated in light at daily alternating temperature regimes of 15°/6°, 20°/10°, and 25°/15°C. Dormancy was not due to impermeability of the stony endocarp surrounding each seed, and embryos did not grow prior to radicle emergence. Thus, seeds did not have physical, morphological, or morphophysiological dormancy. Long periods of either cold stratification (20 or 32 wk) or warm stratification (16 wk) resulted in a maximum of 22-38 and 10% germination, respectively, in light at 25°/15°C. After 12 wk warm stratification plus 20 wk cold stratification, 83-93% of the seeds germinated in light at the three temperature regimes. For a cold stratification period of 20 wk, germination increased with increase in length of the preceding warm stratification treatment. Gibberellic acid (GA(3)) promoted germination of 77-87% of the seeds. Based on dormancy-breaking requirements and response to GA(3), 62-78% of the seeds had intermediate physiological dormancy; the others had nondeep physiological dormancy. Contrary to suggestions of several other investigators that warm stratification is required to make the endocarp permeable to water via its breakdown by microorganisms, our results with E. hermaphroditum show that this is not the case. In this species, warm stratification is part of the dormancy-breaking requirement of embryos in seeds with intermediate physiological dormancy.     

Effects of pre-treatments and temperature on seed viability and germination of Juniperus macrocarpa Sm.

The effects of collecting season, collection site, laboratory pre-treatments and temperatures on seed viability and germination of Juniperus macrocarpa were investigated. Ripe cones were collected in four Sardinian dune systems, in two seasons, from plant and soil. Warm (W) and cold (C) stratifications, two combinations of them (W + C, C + W), and no pre-treatment (0) were applied. Seeds were incubated in a range of constant (10–25 °C) and an alternating (25/10 °C) temperature regime. Seed viability was low (ca. 40%) and varied significantly according to the collecting season. Seed germination was also low (ca. 10%), the 0 and W were the most effective pre-treatments on stimulating germination. The best germination temperature, without any pre-treatment, was 15 °C (ca. 20%). J. macrocarpa seeds are dormant and the achieved results suggested that the presence of secondary dormancy is induced by cold stratification. Spring appeared to be the best season for seed collecting, whereas autumn was the best for sowing. These results give new findings for restoration activities on this species.     

Post-dispersal embryo development, germination phenology, and seed dormancy in Cardiocrinum cordatum var. glehnii (Liliaceae s. str.), a perennial herb of the broadleaved deciduous forest in Japan

In an investigation of seed germination in Cardiocrinum cordatum var. glehnii, embryos in fresh seeds in October were underdeveloped and did not grow until September of the following year. Then, they grew rapidly and had fully elongated by early November. In the second spring after dispersal, radicles emerged under snow in late March and after snowmelt in April. Cotyledons emerged soon after radicles. In several laboratory experiments, embryos grew at 15°/5°C (light 12 h/ dark 12 h) following 25°/15°C. Radicles emerged from seeds with fully elongated embryos at 5°-15°C after cold stratification at 0°-5°C. Cotyledons emerged in 2 wk from seeds with a radicle at 15°/5°C to 30°/20°C. Although seeds require c. 18-19 mo after dispersal to germinate in nature, under controlled conditions, they required only 9 mo with a sequence of 25°/15°C → 15°/5°C → 0°-5°C → 15°/5°C. This is practical knowledge for propagation of plants from seeds. GA(3) treatment partially substituted for the high temperature requirement. Based on dormancy-breaking requirements, the seeds have deep simple morphophysiological dormancy (MPD). A literature review of seed dormancy in taxa of Liliaceae s. str. showed that phylogenetic position in this case is not a good predictor of level of MPD.     

The temperature dimension of the seed germination niche in fen wetlands

The seed germination niche partly determines adaptation, ecological breadth and geographic range in plant species. In temperate wetlands, environmental temperature is the chief regulator of germination timing, but the ecological significance of high and low temperatures during dormancy break and germination is still poorly understood. Our aim was to characterize the temperature dimension of the germination niche in mountain base-rich fens, determining (1) the effect of different temperatures on dormancy break and germination, and (2) whether different germination strategies may be identified at the species level. We conducted laboratory germination experiments with seeds of 15 species from these habitats, collected in 18 fen sites in the Cantabrian Mountains (Spain) for two consecutive years. In all the species, the seeds were totally or conditionally dormant at dispersal and stratification produced a significant increase of germination. In most cases, there was not an obligatory requirement for cold temperatures during dormancy break, since warm stratification promoted germination as well. Although the optimal germination thermoperiod was generally high (30/20 °C), most species could also germinate at lower temperatures after cold-stratification. We also identified a group of species associated to cold-water springs that germinated only at low temperatures. Our results demonstrate that dormancy break in mountain base-rich fens does not obligatorily depend on cold temperatures during overwintering. Furthermore, germination at cool temperatures may be more widespread in wetland habitats than previously thought. The existence of two distinctive germination strategies, ‘warm’ and ‘cool’, can potentially give rise to divergent species responses to climate change.     

Role of endogenous hormones,glumes, endosperm and temperature on germination of Leymus chinensis (Poaceae) seeds during development

Aims Leymus chinensis is an original dominant plant in the Songnen grassland, and it has great value for restoration of severely degraded land. However, seeds are dormant, and low germination percentage is a problem for restoring L.chinensis grassland. The mechanism of seed dormancy is not been well understood. The primary aims of the present study were to investigate the dormancy mechanism of L.chinensis seeds (caryopses) with reference to the role of embryo-covering layers, endogenous hormones and temperature.Methods Changes in concentration of the endogenous hormones GA 3, indoleacetic acid (IAA), zeatin riboside (ZR) and abscisic acid (ABA) in L.chinensis seeds from anthesis to maturity were measured by the enzyme-linked immunosorbent assay method. Germination at different stages of maturity were tested at 16/28°C, 5/28°C and 5/35°C for intact seeds with glumes (control), intact seeds with glumes removed (naked-whole seeds) and intact seeds with glumes and one-half of the endosperm removed (naked-half seeds).Important findings Of the four endogenous hormones monitored, only the concentration of ZR differed significantly between the beginning and the end of seed development (increased); the GA 3 /ABA ratio also did not differ. Rank of germination percentage of control at the three temperature regimens was 5/28°C> 16/28°C> 5/35°C. Germination percentage of the naked-half seeds reached 100% under the three temperature regimens. We concluded that dormancy of L.chinensis seeds is not mainly controlled by endogenous hormones. Germination temperature, mechanical resistance of glumes and inhibition of endosperm are the main factors controlling dormancy and germination of L.chinensis seeds.     

Hormonal and temperature regulation of seed dormancy and germination in <Emphasis Type="Italic">Leymus chinensis</Emphasis>

Fluctuating temperature plays a critical role in determining the timing of seed germination in many plant species. However, the physiological and biochemical mechanisms underlying such a response have been paid little attention. The present study investigated the effect of plant growth regulators and cold stratification in regulating Leymus chinensis seed germination and dormancy response to temperature. Results showed that seed germination was less than 2 % at all constant temperatures while fluctuating temperature significantly increased germination percentage. The highest germination was 71 % at 20/30 °C. Removal of the embryo enclosing material of L. chinensis seed germinated to 74 %, and replaced the requirement for fluctuating temperature to germinate, by increasing embryo growth potential. Applications of GA₄₊₇ significantly increased seed germination at constant temperature. Also, inhibition of GA biosynthesis significantly decreased seed germination at fluctuating temperatures depending upon paclobutrazol concentration. This implied GA was necessary for non-dormant seed germination and played an important role in regulating seed germination response to temperature. Inhibition of ABA biosynthesis during imbibition completely released seed dormancy at 20/30 °C, but showed no effect on seed germination at constant temperature, suggesting ABA biosynthesis was important for seed dormancy maintenance but may not involve in seed germination response to temperature. Cold stratification with water or GA₃ induced seed into secondary dormancy, but this effect was reversed by exogenous FL, suggesting ABA biosynthesis during cold stratification was involved in secondary dormancy. Also, cold stratification with FL entirely replaced the requirement of fluctuating temperature for germination with seeds having 73 % germination at constant temperature. This appears to be attributed to inhibition of ABA biosynthesis and an increase of GA biosynthesis during cold stratification, leading to an increased embryo growth potential. We suggest that fluctuating temperature promotes seed germination by increasing embryo growth potential, mainly attributed to GA biosynthesis during imbibitions. ABA is important for seed dormancy maintenance and induction but showed less effect on non-dormant seed germination response to temperature.     

Environmental regulation of dormancy loss in seeds of Lomatium dissectum (Apiaceae)

Background and Aims

Lomatium dissectum (Apiaceae) is a perennial, herbaceous plant of wide distribution in Western North America. At the time of dispersal, L. dissectum seeds are dormant and have under-developed embryos. The aims of this work were to determine the requirements for dormancy break and germination, to characterize the type of seed dormancy, and to determine the effect of dehydration after embryo growth on seed viability and secondary dormancy.

Methods

The temperature requirements for embryo growth and germination were investigated under growth chamber and field conditions. The effect of GA₃ on embryo growth was also analysed to determine the specific type of seed dormancy. The effect of dehydration on seed viability and induction of secondary dormancy were tested in seeds where embryos had elongated about 4-fold their initial length. Most experiments examining the nature of seed dormancy were conducted with seeds collected at one site in two different years. To characterize the degree of variation in dormancy-breaking requirements among seed populations, the stratification requirements of seeds collected at eight different sites were compared.

Key Results

Embryo growth prior to and during germination occurred at temperatures between 3 and 6 °C and was negligible at stratification temperatures of 0·5 and 9·1 °C. Seeds buried in the field and exposed to natural winter conditions showed similar trends. Interruption of the cold stratification period by 8 weeks of dehydration decreased seed viability by about 30 % and induced secondary dormancy in the remaining viable seeds. Comparison of the cold stratification requirements of different seed populations indicates that seeds collected from moist habitats have longer cold stratification requirements that those from semiarid environments.

Conclusions

Seeds of L. dissectum have deep complex morphophysiological dormancy. The requirements for dormancy break and germination reflect an adaptation to trigger germination in late winter.Key words: Apiaceae, cold stratification, Lomatium dissectum, morphophysiological dormancy, secondary dormancy, seed germination     

Comparative <Emphasis Type="Italic">in vitro</Emphasis> germination ecology of <Emphasis Type="Italic">Calopogon tuberosus</Emphasis> var. <Emphasis Type="Italic">tuberosus</Emphasis> (Orchidaceae) across its geographic range

Seed responses to temperature are often essential to the study of germination ecology, but the ecological role of temperature in orchid seed germination remains uncertain. The response of orchid seeds to cold stratification have been studied, but the exact physiological role remains unclear. No studies exist that compare the effects of either cold stratification or temperature on germination among distant populations of the same species. In two separate experiments, the role of temperature (25, 22/11, 27/15, 29/19, 33/24°C) and chilling at 10°C on in vitro seed germination were investigated using distant populations of Calopogon tuberosus var. tuberosus. Cooler temperatures promoted germination of Michigan seeds; warmer temperatures promoted germination of South Carolina and north central Florida seeds. South Florida seed germination was highest under both warm and cool temperatures. More advanced seedling development generally occurred at higher temperatures with the exception of south Florida seedlings, in which the warmest temperature suppressed development. Fluctuating diurnal temperatures were more beneficial for germination compared to constant temperatures. Cold stratification had a positive effect on germination among all populations, but South Carolina seeds required the longest chilling treatments to obtain maximum germination. Results from the cold stratification experiment indicate that a physiological dormancy is present, but the degree of dormancy varies across the species range. The variable responses among populations may indicate ecotypic differentiation.     

初生休眠解除状态和干燥处理对水曲柳种子萌发的影响

为探究初生休眠解除状态和干燥处理对水曲柳种子萌发的影响，本文以初生休眠的成熟水曲柳种子为材料，研究经不同裸层积（暖温10周+低温8周、暖温12周+低温8周、暖温10周+低温10周、暖温12周+低温10周）和干燥处理（干燥、不干燥）的水曲柳种子在适宜温度和较高温条件下的萌发表现。结果表明，初生休眠解除状态不同的水曲柳种子在不同温度下的萌发表现具有相似的规律，种子的萌发会受到干燥处理的影响。不经干燥处理的种子解除休眠越充分，其萌发能力就越强，但层积处理后的种子若经过干燥处理，则解除休眠越充分（尤其是低温时间越长），种子萌发能力下降越多。水曲柳种子次生休眠（热休眠）的诱导受种子初生休眠解除状态的影响较小，但受干燥处理影响较大。干燥处理会降低水曲柳种子的萌发能力，尤其是较高温条件下的萌发能力，初生休眠解除越充分的种子萌发受干燥处理影响越大。生产中如需对解除休眠的种子干燥处理，选择暖温10周+低温8周的层积方法处理种子效果最好。     

温水浸种对蒙古黄芪种子萌发特性的影响

研究温水浸种破除蒙古黄芪（Astragalus membranaceus Bge.var.mongholicus（Bge.） Hsiao）种子休眠的适宜条件,采用恒温和变温2种浸种方式,测定蒙古黄芪种子在不同时间、不同温度梯度条件下的发芽率、发芽势和发芽指数。结果表明,温水浸种破除蒙古黄芪种子休眠的效果显著,变温浸种效果略优于恒温浸种。不同温度破除休眠效果排序依次为60℃> 70℃> 80℃,其中60℃变温浸种2 min和5 min的发芽率分别为对照的2.87倍和2.31倍。浸种时间、浸种温度均可显著影响蒙古黄芪种子萌发,与对照相比,随着浸种时间增加,发芽率总体呈现出先急速升高后逐渐降低的趋势;发芽指数与浸种温度极显著正相关;浸种温度与发芽势和发芽指数均显著正相关。本研究结果表明温水浸种可以提高种子发芽率,保持种子活力。     

Response of Amaranthus retroflexus L. seeds to gibberellic acid, ethylene and abscisic acid depending on duration of stratification and burial

Freshly harvested, dormant seeds of Amaranthus retroflexus were unable to germinate at 25 and 35 °C. To release their dormancy at the above temperatures, the seeds were stratified at a constant temperature (4 °C) under laboratory conditions or at fluctuating temperatures in soil or by outdoor burial in soil. Fully dormant, or seeds stratified or buried (2006/2007 and 2007/2008) for various periods were treated with exogenous gibberellic acid (GA₃), ethephon and abscisic acid (ABA). Likewise, the effects of these regulators, applied during stratification, on seed germination were determined. The results indicate that A. retroflexus seed dormancy can be released either by stratification or by autumn–winter burial. The effect of GA₃ and ethylene, liberated from ethephon, applied after various periods of stratification or during stratification, depends on dormancy level. GA₃ did not affect or only slightly stimulated the germination of non-stratified, fully dormant seeds at 25 and 35 °C respectively. Ethylene increased germination at both temperatures. Seed response to GA₃ and ethylene at 25 °C was increased when dormancy was partially removed by stratification at constant or fluctuating temperatures or autumn–winter burial. The response to GA₃ and ethylene increased with increasing time of stratification. The presence of GA₃ and ethephon during stratification may stimulate germination at 35 °C. Thus, both GA₃ and ethylene can partially substitute the requirement for stratification or autumn–winter burial. Both hormones may also stimulate germination of secondary dormant seeds, exhumed in September. The response to ABA decreased in parallel with an increasing time of stratification and burial up to May 2007 or March 2008. Endogenous GA_n, ethylene and ABA may be involved in the control of dormancy state and germination of A. retroflexus. It is possible that releasing dormancy by stratification or partial burial is associated with changes in ABA/GA and ethylene balance and/or sensitivity to these hormones.     

皂荚天然群体间种实表型特性及种子萌发的差异分析

对位于湖北京山、湖南城步、贵州兴义、重庆秀山、四川成都、广西桂林、山东费县和甘肃天水的8个皂荚（Gleditsia sinensis Lam．）天然群体间果实和种子表型特性及种子萌发率的差异进行了研究,在此基础上,对种子特性与萌发率的相关性进行了分析。结果表明：不同皂荚群体间荚果的长度、宽度、厚度和质量以及种子长度、宽度、厚度和百粒质量均存在极显著差异（P〈O．01）;在低纬度、高降雨量和干扰程度相对大的区域产生的荚果及种子均较大。在不同温度（昼／夜温度35℃／20℃、30℃／15℃、25℃／10oC和15℃／5℃）条件下,8个皂荚群体间种子萌发率无显著差异,平均萌发率为7．70％;种皮经人为损伤后种子萌发率总体上有所提高且在不同温度条件下差异显著,其中在昼／夜温度15℃／5℃的条件下种子萌发率低于其他处理温度。在不同温度条件下,种子的长度、宽度、厚度和百粒质量与萌发率均呈正相关,但总体上相关性不显著。研究结果说明：不同皂荚群体的荚果和种子特性变异较大,且在降雨量高的地区其果实和种子较大;低温对皂荚种子的萌发有一定抑制作用;种皮损伤处理可解除皂荚种子的物理性休眠,但皂荚种子还可能存在生理休眠现象;一定程度的种子复合休眠可能是皂荚群体适应不同生境的重要生存策略之一。     

Fire and summer temperatures interact to shape seed dormancy thresholds

Background and AimsIn Mediterranean ecosystems, the heat shock of wildfire disrupts physical seed dormancy in many plant species. This triggers germination in the post-fire environment where seedling establishment is optimal due to decreased competition and increased resource availability. However, to maintain the soil seed bank until a fire occurs, the minimum heat capable of breaking seed dormancy (i.e. the lower heat threshold) must be above the maximum temperatures typically observed in the soil during the summer. We therefore hypothesized that summer temperatures have shaped heat requirements for physical dormancy release. Specifically, we predicted that seeds from populations growing under warmer summers will have higher values of the lower heat threshold.MethodsTo evaluate this prediction, we collected seeds from two Cistus species in 31 populations (20 Cistus albidus and 11 Cistus salviifolius) along a climate gradient of summer temperatures on the eastern coast of Spain. For each population, seeds were treated to 10 min heat shocks, from 30 to 120 °C in 5 °C increments (19 treatments), to simulate increasing heat doses from summer to fire-related temperatures. Seeds were then germinated in the lab.Key ResultsFor all populations, maximum germination was observed when applying temperatures associated with fire. Lower heat thresholds varied among populations, with a positive relationship between summer temperatures at seed population origin and the heat dose required to break dormancy.ConclusionsOur results suggest that fire drives maximum dormancy release for successful post-fire germination, while summer temperatures determine lower heat thresholds for ensuring inter-fire seed bank persistence. Significant among-population variation of thresholds also suggests that post-fire seeder species have some potential to modify their dormancy release requirements in response to changing climate.     

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.