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1.  The Success of the Horse-Chestnut Leaf-Miner, Cameraria ohridella, in the UK Revealed with Hypothesis-Led Citizen Science 
PLoS ONE  2014;9(1):e86226.
Citizen science is an increasingly popular way of undertaking research and simultaneously engaging people with science. However, most emphasis of citizen science in environmental science is on long-term monitoring. Here, we demonstrate the opportunities provided by short-term hypothesis-led citizen science. In 2010, we ran the ‘Conker Tree Science’ project, in which over 3500 people in Great Britain provided data at a national scale of an insect (horse-chestnut leaf-mining moth, Cameraria ohridella) undergoing rapid range-expansion. We addressed two hypotheses, and found that (1) the levels of damage caused to leaves of the horse-chestnut tree, Aesculus hippocastanum, and (2) the level of attack by parasitoids of C. ohridella larvae were both greatest where C. ohridella had been present the longest. Specifically there was a rapid rise in leaf damage during the first three years that C. ohridella was present and only a slight rise thereafter, while estimated rates of parasitism (an index of true rates of parasitism) increased from 1.6 to 5.9% when the time C. ohridella had been present in a location increased from 3 to 6 years. We suggest that this increase is due to recruitment of native generalist parasitoids, rather than the adaptation or host-tracking of more specialized parasitoids, as appears to have occurred elsewhere in Europe. Most data collected by participants were accurate, but the counts of parasitoids from participants showed lower concordance with the counts from experts. We statistically modeled this bias and propagated this through our analyses. Bias-corrected estimates of parasitism were lower than those from the raw data, but the trends were similar in magnitude and significance. With appropriate checks for data quality, and statistically correcting for biases where necessary, hypothesis-led citizen science is a potentially powerful tool for carrying out scientific research across large spatial scales while simultaneously engaging many people with science.
PMCID: PMC3899221  PMID: 24465973
2.  Implications of the lack of desiccation tolerance in recalcitrant seeds 
A suite of interacting processes and mechanisms enables tolerance of desiccation and storage (conservation) of orthodox seeds in the dry state. While this is a long-term option under optimized conditions, dry orthodox seeds are not immortal, with life spans having been characterized as short, intermediate and long. Factors facilitating desiccation tolerance are metabolic “switch-off” and intracellular dedifferentiation. Recalcitrant seeds lack these mechanisms, contributing significantly to their desiccation sensitivity. Consequently, recalcitrant seeds, which are shed at high water contents, can be stored only in the short-term, under conditions not allowing dehydration. The periods of such hydrated storage are constrained by germination that occurs without the need for extraneous water, and the proliferation of seed-associated fungi. Cryopreservation is viewed as the only option for long-term conservation of the germplasm of recalcitrant-seeded species. This is not easily achieved, as each of the necessary procedures imposes oxidative damage. Intact recalcitrant seeds cannot be cryopreserved, the common practice being to use excised embryos or embryonic axes as explants. Dehydration is a necessary procedure prior to exposure to cryogenic temperatures, but this is associated with metabolism-linked injury mediated by uncontrolled reactive oxygen species generation and failing anti-oxidant systems. While the extent to which this occurs can be curtailed by maximizing drying rate (flash drying) it cannot be completely obviated. Explant cooling for, and rewarming after, cryostorage must necessarily be rapid, to avoid ice crystallization. The ramifications of desiccation sensitivity are discussed, as are problems involved in cryostorage, particularly those accompanying dehydration and damage consequent upon ice crystallization. While desiccation sensitivity is a “fact” of seed recalcitrance, resolutions of the difficulties involved germplasm conservation are possible as discussed.
PMCID: PMC3837223  PMID: 24319450
cryopreservation; dehydration; desiccation damage; desiccation sensitivity; ice crystallization; recalcitrant seeds; ROS; vitrification
3.  An attempt to control Cameraria ohridella using an attract-and-kill technique 
Journal of Pest Science  2010;84(2):207-212.
This study estimates the efficacy of an attract-and-kill (A&K) technique to control the horse chestnut leaf miner, Cameraria ohridella Deschka and Dimic (Lepidoptera, Gracillariidae), an invasive insect pest of the horse chestnut, Aesculus hippocastanum L. (Hippocastanaceae). The A&K formulation was dispensed as 50 μl droplets of paste-like matrix, containing C. ohridella sex pheromone, (8E,10Z)-tetradeca-8,10-dienal (85% + pure; 0.16% w/w) and a fast acting contact toxicant, pyrocides (94% pure; 6% w/w), applied directly to the bark of the trees. It was tested in 2003 at rates of 30 and 45 droplets/tree at the Ostrobramska site and at rates of 30, 60 and 90 droplets/tree at the Woloska site in Warsaw, Poland, for the first insect generation. A set of untreated plots (0 droplets/tree) was established at each site as well. The treatment efficacy was estimated using two indices: (1) moth catches in pheromone traps and (2) the number of mines per leaf. Trap catches were significantly higher in the untreated plots than in the treated plots regardless of the application rate in all sites. However, there were no significant differences in leaf damage amongst all plots on each site. At the “Lazienki Krolewskie” park the attractiveness of two types of pheromone sources were compared: traps were baited with rubber septum lures or with A&K droplets. The catches of C. ohridella in traps baited with lures were lower than captures in A&K droplet-baited traps, but the difference was not significant. Possible reasons for the low efficacy of the A&K method in management of C. ohridella and reducing leaf damage are discussed.
PMCID: PMC3088814  PMID: 21654920
Cameraria ohridella; Horse chestnut leafminer; Attract-and-kill; Aesculus hippocastanum
4.  Ecological significance of seed desiccation sensitivity in Quercus ilex 
Annals of Botany  2013;111(4):693-701.
Background and Aims
Several widespread tree species of temperate forests, such as species of the genus Quercus, produce recalcitrant (desiccation-sensitive) seeds. However, the ecological significance of seed desiccation sensitivity in temperate regions is largely unknown. Do seeds of such species suffer from drying during the period when they remain on the soil, between shedding in autumn and the return of conditions required for germination in spring?
To test this hypothesis, the Mediterranean holm oak (Quercus ilex) forest was used as a model system. The relationships between the climate in winter, the characteristics of microhabitats, acorn morphological traits, and the water status and viability of seeds after winter were then investigated in 42 woodlands sampled over the entire French distribution of the species.
Key Results
The percentages of germination and normal seedling development were tightly linked to the water content of seeds after the winter period, revealing that in situ desiccation is a major cause of mortality. The homogeneity of seed response to drying suggests that neither intraspecific genetic variation nor environmental conditions had a significant impact on the level of desiccation sensitivity of seeds. In contrast, the water and viability status of seeds at the time of collection were dramatically influenced by cumulative rainfall and maximum temperatures during winter. A significant effect of shade and of the type of soil cover was also evidenced.
The findings establish that seed desiccation sensitivity is a key functional trait which may influence the success of recruitment in temperate recalcitrant seed species. Considering that most models of climate change predict changes in rainfall and temperature in the Mediterranean basin, the present work could help foresee changes in the distribution of Q. ilex and other oak species, and hence plant community alterations.
PMCID: PMC3605958  PMID: 23388882
Quercus ilex; acorn; desiccation sensitivity; holm oak; drought; ecological filtering; germination; Mediterranean climate; recalcitrance; winter rainfall
5.  Histological Examination of Horse Chestnut Infection by Pseudomonas syringae pv. aesculi and Non-Destructive Heat Treatment to Stop Disease Progression 
PLoS ONE  2012;7(7):e39604.
Since its emergence in Northwest Europe as a pathogen that infects trunks and branches of Aesculus spp. (the horse chestnuts) approximately one decade ago, Pseudomonas syringae pv. aesculi has rapidly established itself as major threat to these trees. Infected trees exhibit extensive necrosis of phloem and cambium, which can ultimately lead to dieback. The events after host entry leading to extensive necrosis are not well documented. In this work, the histopathology of this interaction is investigated and heat-treatment is explored as method to eradicate bacteria associated with established infections. The early wound-repair responses of A. hippocastanum, both in absence and presence of P. s. pv. aesculi, included cell wall lignification by a distinct layer of phloem and cortex parenchyma cells. The same cells also deposited suberin lamellae later on, suggesting this layer functions in compartmentalizing healthy from disrupted tissues. However, monitoring bacterial ingress, its construction appeared inadequate to constrain pathogen spread. Microscopic evaluation of bacterial dispersal in situ using immunolabelling and GFP-tagging of P. s. pv. aesculi, revealed two discriminative types of bacterial colonization. The forefront of lesions was found to contain densely packed bacteria, while necrotic areas housed bacterial aggregates with scattered individuals embedded in an extracellular matrix of bacterial origin containing alginate. The endophytic localization and ability of P. s. pv aesculi to create a protective matrix render it poorly accessible for control agents. To circumvent this, a method based on selective bacterial lethality at 39°C was conceived and successfully tested on A. hippocastanum saplings, providing proof of concept for controlling this disease by heat-treatment. This may be applicable for curing other tree cankers, caused by related phytopathogens.
PMCID: PMC3392261  PMID: 22808044
6.  Escin Chemosensitizes Human Pancreatic Cancer Cells and Inhibits the Nuclear Factor-kappaB Signaling Pathway 
Background. There is an urgent need to develop new treatment strategies and drugs for pancreatic cancer that is highly resistant to radio-chemotherapy. Aesculus hippocastanum (the horse chestnut) known in Chinese medicine as a plant with anti-inflammatory, antiedema, antianalgesic, and antipyretic activities. The main active compound of this plant is Escin (C54H84O23). Objective. To evaluate the effect of Escin alone and combined with chemotherapy on pancreatic cancer cell survival and to unravel mechanism(s) of Escin anticancer activity. Methods. Cell survival was measured by XTT colorimetric assay. Synergistic effect of combined therapy was determined by CalcuSyn software. Cell cycle and induction of apoptosis were evaluated by FACS analysis. Expression of NF-κB-related proteins (p65, IκBα, and p-IκBα) and cyclin D was evaluated by western blot analysis. Results. Escin decreased the survival of pancreatic cancer cells with IC50 = 10–20 M. Escin combined with gemcitabine showed only additive effect, while its combination with cisplatin resulted in a significant synergistic cytotoxic effect in Panc-1 cells. High concentrations of Escin induced apoptosis and decreased NF-κB-related proteins and cyclin D expression. Conclusions. Escin decreased pancreatic cancer cell survival, induced apoptosis, and downregulated NF-κB signaling pathway. Moreover, Escin sensitized pancreatic cancer cells to chemotherapy. Further translational research is required.
PMCID: PMC3825277  PMID: 24282639
7.  Physiology, morphology and phenology of seed dormancy break and germination in the endemic Iberian species Narcissus hispanicus (Amaryllidaceae) 
Annals of Botany  2011;107(6):1003-1016.
Background and Aims
Only very few studies have been carried out on seed dormancy/germination in the large monocot genus Narcissus. A primary aim of this study was to determine the kind of seed dormancy in Narcissus hispanicus and relate the dormancy breaking and germination requirements to the field situation.
Embryo growth, radicle emergence and shoot growth were studied by subjecting seeds with and without an emerged radicle to different periods of warm, cold or warm plus cold in natural temperatures outdoors and under controlled laboratory conditions.
Key Results
Mean embryo length in fresh seeds was approx. 1·31 mm, and embryos had to grow to 2·21 mm before radicle emergence. Embryos grew to full size and seeds germinated (radicles emerged) when they were warm stratified for 90 d and then incubated at cool temperatures for 30 d. However, the embryos grew only a little and no seeds germinated when they were incubated at 9/5, 10 or 15/4 °C for 30 d following a moist cold pre-treatment at 5, 9/5 or 10 °C. In the natural habitat of N. hispanicus, seeds are dispersed in late May, the embryo elongates in autumn and radicles emerge (seeds germinate) in early November; however, if the seeds are exposed to low temperatures before embryo growth is completed, they re-enter dormancy (secondary dormancy). The shoot does not emerge until March, after germinated seeds are cold stratified in winter.
Seeds of N. hispanicus have deep simple epicotyl morphophysiological dormancy (MPD), with the dormancy formula C1bB(root) – C3(epicotyl). This is the first study on seeds with simple MPD to show that embryos in advanced stages of growth can re-enter dormancy (secondary dormancy).
PMCID: PMC3080619  PMID: 21335326
Dormancy break; embryo growth; epicotyl morphophysiological dormancy; germination; Narcissus hispanicus; phenology; secondary dormancy; shoot emergence
8.  Do Conditions During Dormancy Influence Germination of Suaeda maritima? 
Annals of Botany  2008;101(9):1319-1327.
Background and Aims
Seeds of annual halophytes such as Suaeda maritima experience fluctuating salinity, hydration, hypoxia and temperature during dormancy. Germination then occurs in one flush of 2–3 weeks after about 5 months of winter dormancy during which time the seeds can remain in saline, often waterlogged soil. The aim of this study was to investigate the effect of simulated natural conditions during dormancy on germination and to compare this with germination following the usual conditions of storing seeds dry. The effects of hydration, salinity, hypoxia and temperature regimes imposed during dormancy on germination were investigated. Also looked at were the effects of seed size on germination and the interaction between salinity during dormancy and salinity at the time of germination.
Various pre-treatments were imposed on samples of seeds that had been stored dry or wet for different periods of time during the 5 months of natural dormancy. Subsequent germination tests were carried out in conditions that simulated those found in the spring when germination occurs naturally. Various salinities were imposed at germination for a test of interaction between storage salinity and salinity at germination.
Key Results
A temperature of about 15 °C was needed for germination and large seeds germinated earlier and better than small seeds. Cold seawater pre-treatment was necessary for good germination; the longer the saline pre-treatment during the natural dormancy period the better the germination. There appeared to be no effect of any specific ion of the seawater pre-treatment on germination and severe hypoxia did not prevent good germination. A short period of freezing stimulated early germination in dry-stored seed. Storage in cold saline or equivalent osmotic medium appeared to inhibit germination during the natural dormancy period and predispose the seed to germinate when the temperature rose and the salinity fell. Seeds that were stored in cold wet conditions germinated better in saline conditions than those stored dry.
The conditions under which seeds of S. maritima are stored affect their subsequent germination. Under natural conditions seeds remain dormant in highly saline, anoxic mud and then germinate when the temperature rises above about 15 °C and the salinity is reduced.
PMCID: PMC2710252  PMID: 18369238
Suaeda maritima; germination; pre-treatment; salinity; temperature
9.  Pre- and Post-harvest Influences on Seed Dormancy Status of an Australian Goodeniaceae species, Goodenia fascicularis 
Annals of Botany  2008;102(1):93-101.
Background and Aims
The period during which seeds develop on the parent plant has been found to affect many seed characteristics, including dormancy, through interactions with the environment. Goodenia fascicularis (Goodeniaceae) seeds were used to investigate whether seeds of an Australian native forb, harvested from different environments and produced at different stages of the reproductive period, differ in dormancy status.
During the reproductive phase, plants were grown ex situ in warm (39/21 °C) or cool (26/13 °C) conditions, with adequate or limited water availability. The physiological dormancy of resulting seeds was measured in terms of the germination response to warm stratification (34/20 °C, 100 % RH, darkness).
Key Results
Plants in the cool environment were tall and had high above-ground biomass, yet yielded fewer seeds over a shorter, later harvest period when compared with plants in the warm environment. Seeds from the cool environment also had higher viability and greater mass, despite a significant proportion (7 % from the cool-wet environment) containing no obvious embryo. In the warm environment, the reproductive phase was accelerated and plants produced more seeds despite being shorter and having lower above-ground biomass than those in the cool environment. Ten weeks of warm stratification alleviated physiological dormancy in seeds from all treatments resulting in 80–100 % germination. Seeds that developed at warm temperatures were less dormant (i.e. germination percentages were higher) than seeds from the cool environment. Water availability had less effect on plant and seed traits than air temperature, although plants with reduced soil moisture were shorter, had lower biomass and produced fewer, less dormant seeds than plants watered regularly.
Goodenia fascicularis seeds are likely to exhibit physiological dormancy regardless of the maternal environment. However, seeds collected from warm, dry environments are likely to be more responsive to warm stratification than seeds from cooler, wetter environments.
PMCID: PMC2712421  PMID: 18430743
Goodenia fascicularis; Goodeniaceae; Australia; physiological dormancy; seeds; temperature; soil moisture; maternal influence; climate
10.  Ecophysiology of Seed Dormancy in the Australian Endemic Species Acanthocarpus preissii (Dasypogonaceae) 
Annals of Botany  2006;98(6):1137-1144.
• Background and Aims Seedlings of Acanthocarpus preissii are needed for coastal sand dune restoration in Western Australia. However, seeds of this Western Australian endemic have proven to be very difficult to germinate. The aims of this study were to define a dormancy-breaking protocol, identify time of suitable conditions for dormancy-break in the field and classify the type of seed dormancy in this species.
• Methods Viability, water-uptake (imbibition) and seed and embryo characteristics were assessed for seeds collected in 2003 and in 2004 from two locations. The effects of GA3, smoke-water, GA3 + smoke-water and warm stratification were tested on seed dormancy-break. In a field study, soil temperature and the moisture content of soil and buried seeds were monitored for 1 year.
• Key Results Viability of fresh seeds was >90 %, and they had a fully developed, curved-linear embryo. Fresh seeds imbibed water readily, with mass increasing approx. 52 % in 4 d. Non-treated fresh seeds and those exposed to 1000 ppm GA3, 1 : 10 (v/v) smoke-water/water or 1000 ppm GA3 + 1 : 10 (v/v) smoke-water/water germinated <8 %. Fresh seeds germinated to >80 % when warm-stratified for at least 7 weeks at 18/33 °C and then moved to 7/18 °C, whereas seeds incubated continuously at 7/18 °C germinated to <20 %.
• Conclusions Seeds of A. preisii have non-deep physiological dormancy that is released by a period of warm stratification. Autumn (March/April) is the most likely time for warm stratification of seeds of this species in the field. This is the first report of the requirement for warm stratification for dormancy release in seeds of an Australian species.
PMCID: PMC2803588  PMID: 17008351
Acanthocarpus preissii; Dasypogonaceae; physiological dormancy; seed dormancy; seed germination; warm stratification
11.  Seed dormancy and ABA signaling 
Plant Signaling & Behavior  2009;4(11):1035-1048.
The seed is an important organ in higher plants, it is an important organ for plant survival and species dispersion. The transition between seed dormancy and germination represents a critical stage in the plant life cycle and it is an important ecological and commercial trait. A dynamic balance of synthesis and catabolism of two antagonistic hormones, abscisic acid (ABA) and giberellins (GAs), controls the equilibrium between seed dormancy and germination. Embryonic ABA plays a central role in induction and maintenance of seed dormancy and also inhibits the transition from embryonic to germination growth. Therefore, the ABA metabolism must be highly regulated at both temporal and spatial levels during phase of dessication tolerance. On the other hand, the ABA levels do not depend exclusively on the seeds because sometimes it becomes a strong sink and imports it from the roots and rhizosphere through the xylem and/or phloem. These events are discussed in depth here. Likewise, the role of some recently characterized genes belonging to seeds of woody species and related to ABA signaling are also included. Finally, although four possible ABA receptors have been reported, not much is known about how they mediate ABA signaling transduction. However, new publications seem to show that almost all these receptors lack several properties to consider them as such.
PMCID: PMC2819511  PMID: 19875942
ABA/GA balance; ABA in woody plants; ABA-receptors; biosynthetic ABA mutants; rhizosphere ABA; seed dormancy
12.  Quantitative description of the effect of stratification on dormancy release of grape seeds in response to various temperatures and water contents 
Journal of Experimental Botany  2009;60(12):3397-3406.
The effect of stratification on dormancy release of grape seeds crossing from the sub- to the supraoptimal range of temperatures and water contents was analysed by modified threshold models. The stratification impacted on dormancy release in three different ways: (i) dormancy was consistently released with prolonged stratification time when stratified at temperatures of <15 °C; (ii) at 15 °C and 20 °C, the stratification effect initially increased, and then decreased with extended time; and (iii) stratification at 25 °C only reduced germinable seeds. These behaviours indicated that stratification could not only release primary dormancy but also induce secondary dormancy in grape seed. The rate of dormancy release changed linearly in two phases, while induction increased exponentially with increasing temperature. The thermal time approaches effectively quantified dormancy release only at suboptimal temperature, but a quantitative method to integrate the occurrence of dormancy release and induction at the same time could describe it well at either sub- or supraoptimal temperatures. The regression with the percentage of germinable seeds versus stratification temperature or water content within both the sub- and supraoptimal range revealed how the optimal temperature (Tso) and water content (Wso) for stratification changed. The Tso moved from 10.6 °C to 5.3 °C with prolonged time, while Wso declined from >0.40 g H2O g DW−1 at 5 °C to ∼0.23 g H2O g DW−1 at 30 °C. Dormancy release in grape seeds can occur across a very wide range of conditions, which has important implications for their ability to adapt to a changeable environment in the wild.
PMCID: PMC2724692  PMID: 19491305
‘Ceiling’ temperature; dormancy induction; optimum temperature; optimum water content; physiological dormancy; thermal time; Vitis
13.  A Race for Survival: Can Bromus tectorum Seeds Escape Pyrenophora semeniperda-caused Mortality by Germinating Quickly? 
Annals of Botany  2007;99(5):907-914.
Background and Aims
Pathogen–seed interactions may involve a race for seed resources, so that seeds that germinate more quickly, mobilizing reserves, will be more likely to escape seed death than slow-germinating seeds. This race-for-survival hypothesis was tested for the North American seed pathogen Pyrenophora semeniperda on seeds of the annual grass Bromus tectorum, an invasive plant in North America. In this species, the seed germination rate varies as a function of dormancy status; dormant seeds germinate slowly if at all, whereas non-dormant seeds germinate quickly.
Three experimental approaches were utilized: (a) artificial inoculations of mature seeds that varied in primary dormancy status and wounding treatment; (b) naturally inoculated undispersed seeds that varied in primary dormancy status; and (c) naturally inoculated seeds from the carry-over seed bank that varied in degree of secondary dormancy, habitat of origin and seed age.
Key Results
In all three approaches, seeds that germinated slowly were usually killed by the pathogen, whereas seeds that germinated quickly frequently escaped. Pyrenophora semeniperda reduced B. tectorum seed banks. Populations in drier habitats sustained 50 times more seed mortality than a population in a mesic habitat. Older carry-over seeds experienced 30 % more mortality than younger seeds.
Given the dramatic levels of seed death and the ability of this pathogen to reduce seed carry-over, it is intriguing to consider whether P. semeniperda could be used to control B. tectorum through direct reduction of its seed bank.
PMCID: PMC2802916  PMID: 17353206
Biocontrol; biotic resistance; cheatgrass; Drechslera campanulata; invasive species; mycoherbicide; pathogen; seed bank; seed-borne
14.  The role of the persistent fruit wall in seed water regulation in Raphanus raphanistrum (Brassicaceae) 
Annals of Botany  2009;105(1):101-108.
Background and Aims
Dry fruits remain around the seeds at dispersal in a number of species, especially the Brassicaceae. Explanations for this vary, but usually involve mechanisms of innate dormancy. We speculate that, instead, a persistent fruit may give additional protection through control of dehydration, to species growing in arid or Mediterranean environments where water is sporadic.
X-rays and weight measurements were used to determine the extent to which Raphanus raphanistrum seeds within mature fruits imbibe water, and germination tests determined the roles of the fruit and seed coat in seed dormancy. Rates of water uptake and desiccation, and seedling emergence were compared with and without the fruit. Finally, germinability of seeds extracted from fruits was determined after various periods of moist conditions followed by a range of dry conditions.
Key Results
Most seeds rapidly take up water within the fruit, but they do not fully imbibe when compared with naked seeds. The seed coat is more important than the dry fruit wall in maintaining seed dormancy. The presence of a dry fruit slows emergence from the soil by up to 6–8 weeks. The fruit slows the rate of desiccation of the seed to a limited extent. The presence of the fruit for a few days during imbibition somehow primes more seeds to germinate than if the fruit is absent; longer moist periods within the pod appear to induce dormancy.
The fruit certainly modifies the seed environment as external conditions change between wet and dry, but not to a great extent. The major role seems to be: (a) the physical restriction of imbibition and germination; and (b) the release and then re-imposition of dormancy within the seed. The ecological significance of the results requires more research under field conditions.
PMCID: PMC2794069  PMID: 19889801
Wild radish; Raphanus raphanistrum; imbibition; desiccation; dry fruit wall; germination; dormancy; X-ray
15.  Comparative Study of Antioxidant Status in Androgenic Embryos of Aesculus hippocastanum and Aesculus flava 
The Scientific World Journal  2014;2014:767392.
In vivo (leaves and seed embryos) and in vitro (androgenic embryos) antioxidant scavenging activity of Aesculus hippocastanum and Aesculus flava medical plants was examined. Here we report antioxidant enzyme activities of superoxide dismutase, catalase, guaiacol peroxidase and glutathione peroxidase, reduced glutathione quantity, flavonoids, soluble protein contents, quantities of malondialdehyde, and •OH radical presence in the investigated plant samples. Total antioxidant capacity of all the samples of A. hippocastanum and A. flava was determined using FRAP, DPPH, and NO• radical scavenger capacity. The leaves of A. flava collected from the botanical garden exhibited stronger antioxidant activity (higher activities of SOD, and higher quantities of GSH, TSH, TPC, and scavenging abilities of DPPH and NO•, and higher FRAP values and lowest quantities of •OH and MDA) than in vitro obtained cultures. However, the leaves of A. flava showed higher antioxidant activity than the leaves of A. hippocastanum, and therefore they have a stronger tolerance of oxidative stress. Androgenic embryos of both species had low amount of antioxidants due to controlled in vitro environmental conditions (T, photoperiod, humidity, nutritive factors, and pathogen-free). Our results confirmed that we found optimal in vitro conditions for producing androgenic embryos of both Aesculus species. Also, we assume that horse chestnut androgenic embryos can be used as an alternative source for large-scale aescin production.
PMCID: PMC3932234  PMID: 24672369
16.  Germination responses to temperature and water potential in Jatropha curcas seeds: a hydrotime model explains the difference between dormancy expression and dormancy induction at different incubation temperatures 
Annals of Botany  2011;109(1):265-273.
Background and Aims
Jatropha curcas is a drought-resistant tree whose seeds are a good source of oil that can be used for producing biodiesel. A successful crop establishment depends on a rapid and uniform germination of the seed. In this work we aimed to characterize the responses of J. curcas seeds to temperature and water availability, using thermal time and hydrotime analysis,
Thermal and hydrotime analysis was performed on germination data obtained from the incubation of seeds at different temperatures and at different water potentials.
Key Results
Base and optimum temperatures were 14·4 and 30 °C, respectively. Approximately 20 % of the seed population displayed absolute dormancy and part of it displayed relative dormancy which was progressively expressed in further fractions when incubation temperatures departed from 25 °C. The thermal time model, but not the hydrotime model, failed to describe adequately final germination percentages at temperatures other than 25 °C. The hydrotime constant, θH, was reduced when the incubation temperature was increased up to 30 °C, the base water potential for 50 % germination,Ψb(50), was less negative at 20 and 30 °C than at 25 °C, indicating either expression or induction of dormancy. At 20 °C this less negative Ψb(50) explained satisfactorily the germination curves obtained at all water potentials, while at 30 °C it had to be corrected towards even less negative values to match observed curves at water potentials below 0. Hence, Ψb(50) appeared to have been further displaced to less negative values as exposure to 30 °C was prolonged by osmoticum. These results suggest expression of dormancy at 20 °C and induction of secondary dormancy above 25 °C. This was confirmed by an experiment showing that inhibition of germination imposed by temperatures higher than 30 °C, but not that imposed at 20 °C, is a permanent effect.
This study revealed (a) the extremely narrow thermal range within which dormancy problems (either through expression or induction of dormancy) may not be encountered; and (b) the high sensitivity displayed by these seeds to water shortage. In addition, this work is the first one in which temperature effects on dormancy expression could be discriminated from those on dormancy induction using a hydrotime analysis.
PMCID: PMC3241573  PMID: 21917817
Jatropha curcas; Euphorbiaceae; seed germination; dormancy; thermal time; hydrotime; water potential
17.  ABA crosstalk with ethylene and nitric oxide in seed dormancy and germination 
Dormancy is an adaptive trait that enables seed germination to coincide with favorable environmental conditions. It has been clearly demonstrated that dormancy is induced by abscisic acid (ABA) during seed development on the mother plant. After seed dispersal, germination is preceded by a decline in ABA in imbibed seeds, which results from ABA catabolism through 8′-hydroxylation. The hormonal balance between ABA and gibberellins (GAs) has been shown to act as an integrator of environmental cues to maintain dormancy or activate germination. The interplay of ABA with other endogenous signals is however less documented. In numerous species, ethylene counteracts ABA signaling pathways and induces germination. In Brassicaceae seeds, ethylene prevents the inhibitory effects of ABA on endosperm cap weakening, thereby facilitating endosperm rupture and radicle emergence. Moreover, enhanced seed dormancy in Arabidopsis ethylene-insensitive mutants results from greater ABA sensitivity. Conversely, ABA limits ethylene action by down-regulating its biosynthesis. Nitric oxide (NO) has been proposed as a common actor in the ABA and ethylene crosstalk in seed. Indeed, convergent evidence indicates that NO is produced rapidly after seed imbibition and promotes germination by inducing the expression of the ABA 8′-hydroxylase gene, CYP707A2, and stimulating ethylene production. The role of NO and other nitrogen-containing compounds, such as nitrate, in seed dormancy breakage and germination stimulation has been reported in several species. This review will describe our current knowledge of ABA crosstalk with ethylene and NO, both volatile compounds that have been shown to counteract ABA action in seeds and to improve dormancy release and germination.
PMCID: PMC3607800  PMID: 23531630
abscisic acid; dormancy; ethylene; germination; hormone; nitric oxide; seed
18.  Stratification Requirements for Seed Dormancy Alleviation in a Wetland Weed 
PLoS ONE  2013;8(9):e71457.
Echinochloaoryzicola(syn.E. phyllopogon) is an exotic weed of California rice paddies that has evolved resistance to multiple herbicides. Elimination of seedlingsthroughcertain weed control methods can limit the spread of this weed, but is contingent on accurate predictions of germination and emergence timing, which are influenced by seed dormancy levels.In summer annuals, dormancy can often be relieved through stratification, a period of prolonged exposure to cold and moist conditions.We used population-based threshold models to quantify the effects of stratification on seed germination of four E. Oryzicola populations at a range of water potential (Ψ) and oxygen levels. We also determined how stratification temperatures, moisture levels and durations contributed to dormancy release. Stratification released dormancy by decreasing base Ψ and hydrotimerequired for germination and by eliminating any germination sensitivity to oxygen. Stratification also increased average germination rates (GR), which were used as a proxy for relative dormancy levels. Alternating temperatures nearly doubled GR in all populations, indicating that seeds could be partially dormant despite achieving high final germination percentages. Stratification at Ψ = 0 MPa increased GR compared to stratification at lower water potentials, demonstrating that Ψ contributed to regulating dormancy release. Maximum GR occurred after 2-4 weeks of stratification at 0 MPa; GR were often more rapid for herbicide-resistant than for herbicide-susceptible seeds, implying greater dormancy in the latter. Manipulation of field conditions to promote dormancy alleviation of E. oryzicola seeds might improve the rate and uniformity of germination for seed bank depletion through seedling weed control. Our results suggest field soil saturation in winter would contribute towards E. oryzicola dormancy release and decrease the time to seedling emergence.
PMCID: PMC3764233  PMID: 24039714
19.  Nitric Oxide Enhances Desiccation Tolerance of Recalcitrant Antiaris toxicaria Seeds via Protein S-Nitrosylation and Carbonylation 
PLoS ONE  2011;6(6):e20714.
The viability of recalcitrant seeds is lost following stress from either drying or freezing. Reactive oxygen species (ROS) resulting from uncontrolled metabolic activity are likely responsible for seed sensitivity to drying. Nitric oxide (NO) and the ascorbate-glutathione cycle can be used for the detoxification of ROS, but their roles in the seed response to desiccation remain poorly understood. Here, we report that desiccation induces rapid accumulation of H2O2, which blocks recalcitrant Antiaris toxicaria seed germination; however, pretreatment with NO increases the activity of antioxidant ascorbate-glutathione pathway enzymes and metabolites, diminishes H2O2 production and assuages the inhibitory effects of desiccation on seed germination. Desiccation increases the protein carbonylation levels and reduces protein S-nitrosylation of these antioxidant enzymes; these effects can be reversed with NO treatment. Antioxidant protein S-nitrosylation levels can be further increased by the application of S-nitrosoglutathione reductase inhibitors, which further enhances NO-induced seed germination rates after desiccation and reduces desiccation-induced H2O2 accumulation. These findings suggest that NO reinforces recalcitrant seed desiccation tolerance by regulating antioxidant enzyme activities to stabilize H2O2 accumulation at an appropriate concentration. During this process, protein carbonylation and S-nitrosylation patterns are used as a specific molecular switch to control antioxidant enzyme activities.
PMCID: PMC3107241  PMID: 21674063
20.  Proteome analysis of Norway maple (Acer platanoides L.) seeds dormancy breaking and germination: influence of abscisic and gibberellic acids 
BMC Plant Biology  2009;9:48.
Seed dormancy is controlled by the physiological or structural properties of a seed and the external conditions. It is induced as part of the genetic program of seed development and maturation. Seeds with deep physiological embryo dormancy can be stimulated to germinate by a variety of treatments including cold stratification. Hormonal imbalance between germination inhibitors (e.g. abscisic acid) and growth promoters (e.g. gibberellins) is the main cause of seed dormancy breaking. Differences in the status of hormones would affect expression of genes required for germination. Proteomics offers the opportunity to examine simultaneous changes and to classify temporal patterns of protein accumulation occurring during seed dormancy breaking and germination. Analysis of the functions of the identified proteins and the related metabolic pathways, in conjunction with the plant hormones implicated in seed dormancy breaking, would expand our knowledge about this process.
A proteomic approach was used to analyse the mechanism of dormancy breaking in Norway maple seeds caused by cold stratification, and the participation of the abscisic (ABA) and gibberellic (GA) acids. Forty-four proteins showing significant changes were identified by mass spectrometry. Of these, eight spots were identified as water-responsive, 18 spots were ABA- and nine GA-responsive and nine spots were regulated by both hormones. The classification of proteins showed that most of the proteins associated with dormancy breaking in water were involved in protein destination. Most of the ABA- and GA-responsive proteins were involved in protein destination and energy metabolism.
In this study, ABA was found to mostly down-regulate proteins whereas GA up-regulated proteins abundance. Most of the changes were observed at the end of stratification in the germinated seeds. This is the most active period of dormancy breaking when seeds pass from the quiescent state to germination. Seed dormancy breaking involves proteins of various processes but the proteasome proteins, S-adenosylmethionine synthetase, glycine-rich RNA binding protein, ABI3-interacting protein 1, EF-2 and adenosylhomocysteinase are of particular importance. The effect of exogenously applied hormones was not a determining factor for total inhibition (ABA) or stimulation (GA) of Norway maple seed dormancy breaking and germination but proteomic data has proven these hormones play a role.
PMCID: PMC2688491  PMID: 19413897
21.  Seed Anatomy and Water Uptake in Relation to Seed Dormancy in Opuntia tomentosa (Cactaceae, Opuntioideae) 
Annals of Botany  2007;99(4):581-592.
Background and Aims
There is considerable confusion in the literature concerning impermeability of seeds with ‘hard’ seed coats, because the ability to take up (imbibe) water has not been tested in most of them. Seeds of Opuntia tomentosa were reported recently to have a water-impermeable seed coat sensu lato (i.e. physical dormancy), in combination with physiological dormancy. However, physical dormancy is not known to occur in Cactaceae. Therefore, the aim of this study was to determine if seeds of O. tomentosa are water-permeable or water-impermeable, i.e. if they have physical dormancy.
The micromorphology of the seed coat and associated structures were characterized by SEM and light microscopy. Permeability of the seed-covering layers was assessed by an increase in mass of seeds on a wet substrate and by dye-tracking and uptake of tritiated water by intact versus scarified seeds.
Key Results
A germination valve and a water channel are formed in the hilum–micropyle region during dehydration and ageing in seeds of O. tomentosa. The funicular envelope undoubtedly plays a role in germination of Opuntia seeds via restriction of water uptake and mechanical resistance to expansion of the embryo. However, seeds do not exhibit any of three features characteristic of those with physical dormancy. Thus, they do not have a water-impermeable layer(s) of palisade cells (macrosclereids) or a water gap sensu stricto and they imbibe water without the seed coat being disrupted.
Although dormancy in seeds of this species can be broken by scarification, they have physiological dormancy only. Further, based on information in the literature, it is concluded that it is unlikely that any species of Opuntia has physical dormancy. This is the first integrative study of the anatomy, dynamics of water uptake and dormancy in seeds of Cactaceae subfamily Opuntioideae.
PMCID: PMC2802935  PMID: 17298989
México Valley; Opuntia tomentosa; physical dormancy, physiological dormancy; seed anatomy; seed germination; water uptake by seeds
22.  Effect of Temperature, Light and Salinity on Seed Germination and Radicle Growth of the Geographically Widespread Halophyte Shrub Halocnemum strobilaceum 
Annals of Botany  2007;101(2):293-299.
Background and Aims
The small leafy succulent shrub Halocnemum strobilaceum occurs in saline habitats from northern Africa and Mediterranean Europe to western Asia, and it is a dominant species in salt deserts such as those of north-west China. The effects of temperature, light/darkness and NaCl salinity were tested on seed germination, and the effects of salinity were tested on seed germination recovery, radicle growth and radicle elongation recovery, using seeds from north-west China; the results were compared with those previously reported on this species from ‘salt steppes’ in the Mediterranean region of Spain.
Seed germination was tested over a range of temperatures in light and in darkness and over a range of salinities at 25 °C in the light. Seeds that did not germinate in the NaCl solutions were tested for germination in deionized water. Seeds from which radicles had barely emerged in deionized water were transferred to NaCl solutions for 10 d and then back to deionized water for 10 d to test for radicle growth and recovery.
Key Results
Seeds germinated to higher percentages in light than in darkness and at high than at low temperatures. Germination percentages decreased with an increase in salinity from 0·1 to 0·75 m NaCl. Seeds that did not germinate in NaCl solutions did so after transfer to deionized water. Radicle elongation was increased by low salinity, and then it decreased with an increase in salinity, being completely inhibited by ≥2·0 m NaCl. Elongation of radicles from salt solutions <3·0 m resumed after seedlings were transferred to deionized water.
The seed and early seedling growth stages of the life cycle of H. strobilaceum are very salt tolerant, and their physiological responses differ somewhat between the Mediterranean ‘salt steppe’ of Spain and the inland cold salt desert of north-west China.
PMCID: PMC2711011  PMID: 17428834
Halocnemum strobilaceum; halophyte; inland salt desert; radicle growth; radicle growth recovery; seed germination; seed germination recovery
23.  Role of HY5 in abscisic acid response in seeds and seedlings 
Plant Signaling & Behavior  2008;3(11):986-988.
Seed dormancy offers plants an adaptive advantage that is crucial to their successful colonization of land. The decision to make the transition from a dormant seed to a photoautotrophic seedling is a result of a complex interaction of internal hormonal signals and external stimuli such as water, temperature and light. We recently showed that HY5, a well-characterized component in the light signaling pathway, also mediates abscisic acid (ABA) response during seed germination, early seedling growth and root development in Arabidopsis. We proposed that HY5 regulates these ABA responses partly by directly activating the transcription factor gene ABI5. By analyzing the premature germination of hy5 and abi5 single and double mutants, here we demonstrated that HY5 also positively controls seed maturation and dormancy, likely through direct activation of the ABI5 gene. The contrasting role of light regulation of seed development and germination may be important for the adaptation of plants to the environment.
PMCID: PMC2633751  PMID: 19704428
seed dormancy; light; ABA; HY5; ABI5
24.  Seed dormancy in alpine species 
Flora  2011;206(10-3):845-856.
In alpine species the classification of the various mechanisms underlying seed dormancy has been rather questionable and controversial. Thus, we investigated 28 alpine species to evaluate the prevailing types of dormancy. Embryo type and water impermeability of seed coats gave an indication of the potential seed dormancy class. To ascertain the actual dormancy class and level, we performed germination experiments comparing the behavior of seeds without storage, after cold-dry storage, after cold-wet storage, and scarification. We also tested the light requirement for germination in some species. Germination behavior was characterized using the final germination percentage and the mean germination time. Considering the effects of the pretreatments, a refined classification of the prevailing dormancy types was constructed based on the results of our pretreatments. Only two out of the 28 species that we evaluated had predominantly non-dormant seeds. Physiological dormancy was prevalent in 20 species, with deep physiological dormancy being the most abundant, followed by non-deep and intermediate physiological dormancy. Seeds of four species with underdeveloped embryos were assigned to the morphophysiologial dormancy class. An impermeable seed coat was identified in two species, with no additional physiological germination block. We defined these species as having physical dormancy. Light promoted the germination of seeds without storage in all but one species with physiological dormancy. In species with physical dormancy, light responses were of minor importance. We discuss our new classification in the context of former germination studies and draw implications for the timing of germination in the field.
PMCID: PMC3886365  PMID: 24415831
CDSfresh, cold-dry storage of seeds before incubation under long-day conditions; CDSsc, scarification of seeds following cold-dry storage before incubation under long-day conditions; CWSfresh, cold-wet storage of seeds before incubation under long-day conditions; CWSsubs, cold-wet storage subsequent to a germination experiment before incubation under long-day conditions; FGP, final germination percentage; FRESHdark, seeds without storage incubated in darkness; FRESHsc, scarification of seeds without storage before incubation under long-day conditions; FRESHLD, seeds without storage incubated under long-day conditions; GA3, gibberellic acid; MD, morphological dormancy; MGT, mean germination time; MPD, morphophysiological dormancy; ND, non-dormant; PD, physiological dormancy; PY, physical dormancy; PY + PD, combinational dormancy of PY and PD; Cold-dry seed storage; Cold-wet seed storage; Dormancy classification; Embryo morphology; Light response; Scarification
25.  Comparison of germination and seed bank dynamics of dimorphic seeds of the cold desert halophyte Suaeda corniculata subsp. mongolica 
Annals of Botany  2012;110(8):1545-1558.
Background and Aims
Differences in dormancy and germination requirements have been documented in heteromorphic seeds of many species, but it is unknown how this difference contributes to maintenance and regeneration of populations. The primary aim of this study was to compare the seed bank dynamics, including dormancy cycling, of the two seed morphs (black and brown) of the cold desert halophyte Suaeda corniculata and, if differences were found, to determine their influence on regeneration of the species.
Seeds of the two seed morphs were buried, exhumed and tested monthly for 24 months over a range of temperatures and salinities, and germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were also investigated for the two morphs.
Key Results
Black seeds had an annual dormancy/non-dormancy cycle, while brown seeds, which were non-dormant at maturity, remained non-dormant. Black seeds also exhibited an annual cycle in sensitivity of germination to salinity. Seedlings derived from black seeds emerged in July and August and those from brown seeds in May. Seedlings were recruited from 2·6 % of the black seeds and from 2·8 % of the brown seeds in the soil, and only 0·5 % and 0·4 % of the total number of black and brown seeds in the soil, respectively, gave rise to seedlings that survived to produce seeds. Salinity and water stress induced dormancy in black seeds and decreased viability of brown seeds. Brown seeds formed only a transient soil seed bank and black seeds a persistent seed bank.
The presence of a dormancy cycle in black but not in brown seeds of S. corniculata and differences in germination requirements of the two morphs cause them to differ in their germination dynamics. The study contributes to our limited knowledge of dormancy cycling and seed bank formation in species producing heteromorphic seeds.
PMCID: PMC3503492  PMID: 22975287
Dormancy; halophyte seeds; salinity; seed germination; seedling recruitment; seed bank dynamics; Suaeda corniculata

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