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Argyreia is the most recently evolved genus in the Convolvulaceae, and available information suggests that most species in this family produce seeds with physical dormancy (PY). Our aim was to understand the evolution of seed dormancy in this family via an investigation of dormancy, storage behaviour, morphology and anatomy of seeds of five Argyreia species from Sri Lanka. Imbibition, germination and dye tracking of fresh intact and manually scarified seeds were studied. Scanning electron micrographs and hand sections of the hilar area and the seed coat away from the hilar area were compared. Scarified and intact seeds of A. kleiniana , A. hirsuta and A. zeylanica imbibed water and germinated to a high percentage, but only scarified seeds of A. nervosa and A. osyrensis did so. Thus, seeds of the three former species are non-dormant (ND), while those of the latter two have physical dormancy (PY); this result was confirmed by dye-tracking experiments. Since >90% of A. kleiniana, A. hirsuta and A. zeylanica seeds survived desiccation to 10% moisture content (MC) and >90% of A. nervosa and A. osyrensis seeds with a dispersal MC of ~12% were viable, seeds of the five species were desiccation-tolerant. A. nervosa and A. osyrensis have a wide geographical distribution and PY, while A. kleiniana , A. hirsuta and A. zeylanica have a restricted distribution and ND. Although seeds of A. kleiniana are ND, their seed coat anatomy is similar to that of A. osyrensis with PY. These observations suggest that the ND of A. kleiniana , A. hirsuta and A. zeylanica seeds is the result of an evolutionary reversal from PY and that ND may be an adaptation of these species to the environmental conditions of their wet aseasonal habitats. [ABSTRACT FROM AUTHOR]

Aims In dry tropical forests, herbaceous species may have dormancy mechanisms and form persistent and transient seed banks in the soil. Evolutionarily acquired, these mechanisms are efficient for the establishment and survival of these herbs, especially in forests with unpredictable climates, such as the Caatinga. Thus, our objective was to verify whether the studied herbaceous species adopt the physical dormancy mechanism and how these natural barriers are overcome, to understand the temporal dynamics existing in the soil seed bank from a Brazilian dry tropical forest. Methods Seeds of five native herbaceous species from the Caatinga forests were selected and submitted to pre-germinative treatments for verifying the presence of physical dormancy. We collected soil samples in the rainy and dry seasons for four consecutive years and monitored the emergence of the selected herbaceous in the greenhouse. We verified the differences in germination and seed bank emergence in the soil by generalized linear models. Important Findings The presence and absence of physical dormancy were observed in seeds from Caatinga herbaceous species. We found intraspecific and interspecific differences in the herbaceous emergence from soil seed banks between years and climatic seasons. In perennial herbs, consecutive lack of emergence between seasons and years was frequent, which suggests a direct relationship with the mechanism of physical dormancy and the environmental conditions necessary to overcome integument barriers. In these species, seed dimorphism and dormancy may confer additional advantages to their survival. Moreover, presenting intermediate levels of physical dormancy in an annual species may be an evolutionary adjustment to rainfall unpredictability. In contrast, we found that the annual herb without dormancy is more sensitive to seasonal and interannual climate changes, as evidenced by the increase and significant reduction of its emergence in the soil seed bank. These differences acquired evolutionarily are advantageous for the establishment of herbaceous populations, mainly in semiarid regions with an unpredictable climate. [ABSTRACT FROM AUTHOR]

Background and Aims In many flammable ecosystems, physically dormant seeds show dormancy-break patterns tied to fire, but the link between heat shock and germination in the tropical savannas of Africa and South America remains controversial. Seed heat tolerance is important, preventing seed mortality during fire passage, and is usually predicted by seed traits. This study investigated the role of fire frequency (ecological effects) and seed traits through phylogenetic comparison (historical effects), in determining post-fire germination and seed mortality in legume species of the Cerrado, a tropical savanna–forest mosaic. Methods Seeds of 46 legume species were collected from three vegetation types (grassy savannas, woody savannas and forests) with different fire frequencies. Heat shock experiments (100 °C for 1 min; 100 °C for 3 min; 200 °C for 1 min) were then performed, followed by germination and seed viability tests. Principal component analysis, generalized linear mixed models and phylogenetic comparisons were used in data analyses. Key Results Heat shocks had little effect on germination, but seed mortality was variable across treatments and species. Seed mortality was lowest under the 100 °C 1 min treatment, and significantly higher under 100 °C 3 min and 200 °C 1 min; larger seed mass decreased seed mortality, especially at 200 °C. Tree species in Detarioideae had the largest seeds and were unaffected by heat. Small-seeded species (mostly shrubs from grassy savannas) were relatively sensitive to the hottest treatment. Nevertheless, the presence of physical dormancy helped to avoid seed mortality in small-seeded species under the hottest treatment. Conclusions Physical dormancy-break is not tied to fire in the Cerrado mosaic. Heat tolerance appears in both forest and savanna species and is predicted by seed traits (seed mass and physical dormancy), which might have helped forest lineages to colonize the savannas. The results show seed fire responses are better explained by historical than ecological factors in the Cerrado, contrasting with different fire-prone ecosystems throughout the world. [ABSTRACT FROM AUTHOR]

Background A water-impermeable testa acts as a barrier to a seed's imbibition, thereby imposing dormancy. The physical and functional properties of the macrosclereids are thought to be critical determinants of dormancy; however, the mechanisms underlying the maintenance of and release from dormancy in pea are not well understood. Methods Seeds of six pea accessions of contrasting dormancy type were tested for their ability to imbibe and the permeability of their testa was evaluated. Release from dormancy was monitored following temperature oscillation, lipid removal and drying. Histochemical and microscopic approaches were used to characterize the structure of the testa. Key results The strophiole was identified as representing the major site for the entry of water into non-dormant seeds, while water entry into dormant seeds was distributed rather than localized. The major barrier for water uptake in dormant seeds was the upper section of the macrosclereids, referred to as the 'light line'. Dormancy could be released by thermocycling, dehydration or chloroform treatment. Assays based on either periodic acid or ruthenium red were used to visualize penetration through the testa. Lipids were detected within a subcuticular waxy layer in both dormant and non-dormant seeds. The waxy layer and the light line both formed at the same time as the establishment of secondary cell walls at the tip of the macrosclereids. Conclusions The light line was identified as the major barrier to water penetration in dormant seeds. Its outer border abuts a waxy subcuticular layer, which is consistent with the suggestion that the light line represents the interface between two distinct environments – the waxy subcuticular layer and the cellulose-rich secondary cell wall. The mechanistic basis of dormancy break includes changes in the testa's lipid layer, along with the mechanical disruption induced by oscillation in temperature and by a decreased moisture content of the embryo. [ABSTRACT FROM AUTHOR]

Background and Aims Different structures have been shown to act as a water gap in seeds with physical dormancy (PY), and in Fabaceae they are commonly located in the hilar region. However, the function of the pleurogram, a structure in the extra-hilar region that is common in legume seeds, remains unknown. Our aims were to review the literature for occurrence of the pleurogram in Fabaceae, determine if the pleurogram can open, and compare the functional morpho-anatomy of water gaps in seeds of 11 Senna species. Methods Imbibition tests showed that all 11 species had PY. Structural features of the hilar and extra-hilar regions of the seeds were investigated using light and scanning electron microscopy, and dye-tracking was performed to trace the pathways of water through the seed coat. Key Results A pleurogram has been reported for 37 legume genera. Water gaps differed among Senna species, with lens, hilum, micropyle and pleurogram taking up water after PY was broken. In Senna alata seeds, only the pleurogram acted as a water gap, whereas in S. reniformis and S. silvestris water entered the seed through both the pleurogram and the hilar region. In the pleurogram of S. alata and S. reniformis, the palisade layer moved outward, exposing the hourglass cells, whereas in S. silvestris the palisade layer was broken. Conclusions The pleurogram acts as a water gap in some of the 11 Senna species examined, but it is non-functional in others. Opening the pleurogram occurs due to the formation of a linear slit in the palisade layer. The pleurogram is of functional significance by creating a wide opening, whereby water can reach the embryo and start germination. This is the first report of the pleurogram functioning as a water gap. Because this structure is shared by at least 37 genera, it also may be a water gap in many other legume species. [ABSTRACT FROM AUTHOR]

Aims Seed dormancy and the soil seed bank are crucial to plant regeneration strategy, especially in semiarid ecosystems with unpredictable precipitation. The aim of this study was to investigate how seed dormancy is controlled by environmental factors and how it is correlated with the soil seed bank and regeneration of the perennial legume Oxytropis racemosa, a dominant perennial herb in Mu Us Sandland of semiarid China. Methods Germination and imbibition experiments on fresh intact and scarified seeds of O. racemosa were used to identify physical dormancy (PY) in seeds of this species. Soil seed bank dynamics, timing of seedling emergence and the fate of buried seeds in the natural habitat were investigated. Important Findings PY was broken by mechanical scarification or wet heat/ice water cycles but not solely by dry heat or wet heat treatment. The soil seed bank exhibited seasonal changes in the number of seeds, which was highest in September and lowest in July. Seeds buried at different sand depths gradually lost dormancy; 20-42% of the seeds remained dormant after 20 months of burial. Dormancy break occurs gradually throughout the year. Our results indicate that O. racemosa exhibits hardcoatedness heterogeneity that spreads germination of a seed cohort between seasons and years in the semiarid environment, where the amount of precipitation during the growing season is highly variable. [ABSTRACT FROM AUTHOR]

In physically dormant species, fire-generated heat breaks seed dormancy, with dormancy-breaking temperature thresholds varying among species. Fire-related heating decreases with depth, due to the insulating effects of soil. Concurrently, smaller-seeded species are restricted to germinating near the surface because of limited reserves within such seeds. We hypothesized that dormancy-breaking temperature thresholds of physically dormant species would be higher in smaller-seeded species, to ensure emergence is restricted to shallower soil depths, and that seed survivorship would follow the same pattern. This was tested experimentally for 14 species from south-eastern Australia, and by using a larger data set of species compiled from the literature to assess if any patterns hold across a broader group. Seed size was negatively related to dormancy-breaking temperature thresholds. Mortality at 100 °C showed a positive relationship with seed size. Our findings suggest that small-seeded species are subject to firerelated selection pressure that results in higher dormancy-breaking temperature thresholds and resistant to hotter temperatures, which may act as a depth detection mechanism. By using a broader range of species, we highlight that this relationship is strong and representative of species across a number of phylogenetic groups. [ABSTRACT FROM AUTHOR]

Aims The study of the adaptive potential of the germination patterns of invading species enables us to identify some traits linked with their capacity to colonize new sites, and to gain a better understanding of their area of distribution. The aim of this study is to determine the germination pattern of Ulex europaeus, a cosmopolitan invasive species, in relation to temperature and to explore its potential evolution and invasiveness in a tropical region, following its introduction from Europe. Methods We studied the germination pattern of U. europaeus (the common gorse) to test both physical dormancy and germination capacity within the range of temperatures found in the native and invasive regions. To understand its germination pattern and its evolution, the rate and the speed of germination, as well as the percentage of seeds that became mouldy during the experiment, have been compared between a native habitat, France and a habitat into which it has been introduced, the tropical island of La Reunion. Important Findings The results show that gorse seeds germinate in large quantities, possess the ability to germinate under a wide range of temperatures and they confirm the physical dormancy of the seeds (caused by seed coat impermeability). The decrease in germination from 25°C upwards, coupled with an increase in the rate of moulding help to explain its restricted distribution at altitude in tropical environments. For scarified seeds, we have not detected any difference between the two regions, neither in the percentage of germinated seeds, nor in the percentage of mouldy seeds. However, seeds from Reunion germinate faster at 20°C than seeds from France and a greater number of seeds from Reunion are able to germinate without scarification (10-60% for Reunion versus 0-10% for France). These results suggest that while preserving the advantages of the native habitat, in Reunion gorse develops a strategy which favours the rapid occupation of new sites. [ABSTRACT FROM AUTHOR]

Background and aims Seed dormancy enhances fitness by preventing seeds from germinating when the probability of seedling survival and recruitment is low. The onset of physical dormancy is sensitive to humidity during ripening; however, the implications of this mechanism for seed bank dynamics have not been quantified. This study proposes a model that describes how humidity-regulated dormancy onset may control the accumulation of a dormant seed bank, and seed experiments are conducted to calibrate the model for an Australian Fabaceae, Acacia saligna. The model is used to investigate the impact of climate on seed dormancy and to forecast the ecological implications of human-induced climate change. Methods The relationship between relative humidity and dormancy onset was quantified under laboratory conditions by exposing freshly matured non-dormant seeds to constant humidity levels for fixed durations. The model was field-calibrated by measuring the response of seeds exposed to naturally fluctuating humidity. The model was applied to 3-hourly records of humidity spanning the period 1972–2007 in order to estimate both temporal variability in dormancy and spatial variability attributable to climatic differences among populations. Climate change models were used to project future changes in dormancy onset. Key Results A sigmoidal relationship exists between dormancy and humidity under both laboratory and field conditions. Seeds ripened under field conditions became dormant following very short exposure to low humidity (<20 %). Prolonged exposure at higher humidity did not increase dormancy significantly. It is predicted that populations growing in a temperate climate produce 33–55 % fewer dormant seeds than those in a Mediterranean climate; however, dormancy in temperate populations is predicted to increase as a result of climate change. Conclusions Humidity-regulated dormancy onset may explain observed variation in physical dormancy. The model offers a systematic approach to modelling this variation in population studies. Forecast changes in climate have the potential to alter the seed bank dynamics of species with physical dormancy regulated by this mechanism, with implications for their capacity to delay germination and exploit windows for recruitment. [ABSTRACT FROM AUTHOR]

Background and Aims Physical dormancy (PY) occurs in seeds or fruits of 18 angiosperm families and is caused by a water-impermeable palisade cell layer(s) in seed or fruit coats. Prior to germination, the seed or fruit coat of species with PY must become permeable in order to imbibe water. Breaking of PY involves formation of a small opening(s) (water gap) in a morpho-anatomically specialized area in seeds or fruits known as the water-gap complex. Twelve different water-gap regions in seven families have previously been characterized. However, the water-gap regions had not been characterized in Cucurbitaceae; clade Cladrastis of Fabaceae; subfamilies Bombacoideae, Brownlowioideae and Bythnerioideae of Malvaceae; Nelumbonaceae; subfamily Sapindoideae of Sapindaceae; Rhamnaceae; or Surianaceae. The primary aims of this study were to identify and describe the water gaps of these taxa and to classify all the known water-gap regions based on their morpho-anatomical features. Methods Physical dormancy in 15 species was broken by exposing seeds or fruits to wet or dry heat under laboratory conditions. Water-gap regions of fruits and seeds were identified and characterized by use of microtome sectioning, light microscopy, scanning electron microscopy, dye tracking and blocking experiments. Key Results Ten new water-gap regions were identified in seven different families, and two previously hypothesized regions were confirmed. Water-gap complexes consist of (1) an opening that forms after PY is broken; (2) a specialized structure that occludes the gap; and (3) associated specialized tissues. In some species, more than one opening is involved in the initial imbibition of water. Conclusions Based on morpho-anatomical features, three basic water-gap complexes (Types-I, -II and -III) were identified in species with PY in 16 families. Depending on the number of openings involved in initial imbibition, the water-gap complexes were sub-divided into simple and compound. The proposed classification system enables understanding of the relationships between the water-gap complexes of taxonomically unrelated species with PY. [ABSTRACT FROM PUBLISHER]

Background and Aims The involvement of two steps in the physical dormancy (PY)-breaking process previously has been demonstrated in seeds of Fabaceae and Convolvulaceae. Even though there is a claim for a moisture-controlled stepwise PY-breaking in some species of Geraniaceae, no study has evaluated the role of temperature in the PY-breaking process in this family. The aim of this study was to determine whether a temperature-controlled stepwise PY-breaking process occurs in seeds of the winter annuals Geranium carolinianum and G. dissectum. Methods Seeds of G. carolinianum and G. dissectum were stored under different temperature regimes to test the effect of storage temperature on PY-break. The role of temperature and moisture regimes in regulating PY-break was investigated by treatments simulating natural conditions. Greenhouse (non-heated) experiments on seed germination and burial experiments (outdoors) were carried out to determine the PY-breaking behaviour in the natural habitat. Key Results Irrespective of moisture conditions, sensitivity to the PY-breaking step in seeds of G. carolinianum was induced at temperatures ≥20 °C, and exposure to temperatures ≤20 °C made the sensitive seeds permeable. Sensitivity of seeds increased with time. In G. dissectum, PY-break occurred at temperatures ≥20 °C in a single step under constant wet or dry conditions and in two steps under alternate wet–dry conditions if seeds were initially kept wet. Conclusions Timing of seed germination with the onset of autumn can be explained by PY-breaking processes involving (a) two temperature-dependent steps in G. carolinianum and (b) one or two moisture-dependent step(s) along with the inability to germinate under high temperatures in G. dissectum. Geraniaceae is the third of 18 families with PY in which a two-step PY-breaking process has been demonstrated. [ABSTRACT FROM AUTHOR]

Background and Aims The ‘hinged valve gap’ has been previously identified as the initial site of water entry (i.e. water gap) in physically dormant (PY) seeds of Geranium carolinianum (Geraniaceae). However, neither the ontogeny of the hinged valve gap nor acquisition of PY by seeds of Geraniaceae has been studied previously. The aims of the present study were to investigate the physiological events related to acquisition of PY and the ontogeny of the hinged valve gap and seed coat of G. carolinianum. Methods Seeds of G. carolinianum were studied from the ovule stage until dispersal. The developmental stages of acquisition of germinability, physiological maturity and PY were determined by seed measurement, germination and imbibition experiments using intact seeds and isolated embryos of both fresh and slow-dried seeds. Ontogeny of the seed coat and water gap was studied using light microscopy. Key Results Developing seeds achieved germinability, physiological maturity and PY on days 9, 14 and 20 after pollination (DAP), respectively. The critical moisture content of seeds on acquisition of PY was 11 %. Slow-drying caused the stage of acquisition of PY to shift from 20 to 13 DAP. Greater extent of cell division and differentiation at the micropyle, water gap and chalaza than at the rest of the seed coat resulted in particular anatomical features. Palisade and subpalisade cells of varying forms developed in these sites. A clear demarcation between the water gap and micropyle is not evident due to their close proximity. Conclusions Acquisition of PY in seeds of G. carolinianum occurs after physiological maturity and is triggered by maturation drying. The micropyle and water gap cannot be considered as two separate entities, and thus it is more appropriate to consider them together as a ‘micropyle–water-gap complex’. [ABSTRACT FROM AUTHOR]

Background and Aims: Physical dormancy in seeds of species of Geraniaceae is caused by a water-impermeable palisade layer in the outer integument of the seed coat and a closed chalaza. The chalazal cleft has been reported to be the water gap (i.e. location of initial water entry) in innately permeable seeds of Geraniaceae. The primary aim of this study was to re-evaluate the location of the water gap and to characterize its morphology and anatomy in physically dormant seeds of Geraniaceae, with particular reference to G. carolinianum. [ABSTRACT FROM PUBLISHER]

Background and Aims: The Sapindaceae is one of 17 plant families in which seed dormancy is caused by a water-impermeable seed or fruit coat (physical dormancy, PY). However, until now the water gap in Sapindaceae had not been identified. The primary aim of this study was to identify the water gap in Dodonaea petiolaris (Sapindaceae) seeds and to describe its basic morphology and anatomy. [ABSTRACT FROM PUBLISHER]

• Background and Aims Hard-seeded (physical) dormancy is common among plants, yet mechanisms for dormancy release are poorly understood, especially in the tropics. The following questions are asked: (a) whether dormancy release in seed banks of the tropical shrub Parkinsonia aculeata (Caesalpiniaceae) is determined by wet heat (incubation under wet, warm to hot, conditions); and (b) whether its effect is modified by microclimate.• Methods A seed burial trial was conducted in the wet–dry tropics (northern Australia) to compare dormancy release across different habitats (open, artificial cover, ground cover and canopy cover), burial depths (0, 3 and 20 cm) and burial durations (2, 6 and 14 weeks). Results were compared with predictions using a laboratory-derived relationship between wet heat and dormancy release, and microclimate data collected during the trial.• Key Results Wet heat (defined as the soil temperature above which seeds were exposed to field capacity or higher for a cumulative total of 24 h) was 43·6 °C in the 0 cm open treatment, and decreased with increasing shade and depth to 29·5 °C at 20 cm under canopy cover. The dormancy release model showed that wet heat was a good predictor of the proportion of seeds remaining dormant. Furthermore, dormancy release was particularly sensitive to wet heat across the temperature range encountered across treatments. This resulted in a 16-fold difference in dormancy levels between open (<5 % of seeds still dormant) and covered (82 %) microhabitats.• Conclusions These results demonstrate that wet heat is the principal dormancy release mechanism for P. aculeata when conditions are hot and wet. They also highlight the potential importance of microclimate in driving the population dynamics of such species. [ABSTRACT FROM AUTHOR]

The preservation of plant species under ex situ conditions in seed banks strongly depends on seed longevity. However, detailed knowledge on this seed ecological aspect is limited and comparative studies from central European habitats are scarce. Therefore, we investigated the seed longevity of 39 calcareous grassland species in order to assess the prospects of ex situ storage of seeds originating from a single, strongly threatened habitat. Seed longevity (p 50 ) was determined by artificially ageing the seeds under rapid ageing conditions (45 °C and 60 % eRH (equilibrium relative humidity)), testing for germination and calculating survival curves. We consulted seed and germination traits that are expected to be related to seed longevity. P 50 values strongly varied within calcareous grassland species. The p 50 values ranged between 3.4 and 282.2 days. We discovered significantly positive effects of physical dormancy and endosperm absence on p 50 . Physiological dormancy was associated to comparatively short longevity. These relationships remained significant when accounting for phylogenetic effects. Seed mass, seed shape, and seed coat thickness were not associated with longevity. We therefore recommend more frequent viability assessments of stored endospermic, non-physically and physiologically dormant seeds. [ABSTRACT FROM AUTHOR]