Polyploidy affects the seed,dormancy and seedling characteristics of a perennial grass,conferring an advantage in stressful climates

• Polyploidy (the state of having more than two genome copies) is widely distributed in flowering plants and can vary within species, with polyploid races often associated with broad ecological tolerances. Polyploidy may influence within‐species variation in seed development, germination and establishment. We hypothesized that interactions between polyploidy and the seed developmental environment would affect subsequent dormancy, germination and early growth traits, particularly in stressful environments.
• Using seeds developed in a common garden under ambient and warmed conditions, we conducted germination trials under drought and temperature stress, and monitored the subsequent growth of seedlings. The study species, Themeda triandra, is a widespread, keystone, Australian native grass and a known polyploid complex.
• Tetraploid plants produced heavier, more viable seeds than diploids. Tetraploids were significantly more dormant than diploids, regardless of seed developmental environment. Non‐dormant tetraploids were more sensitive to germination stress compared to non‐dormant diploids. Finally, tetraploid seedlings were larger and grew faster than diploids, usually when maternal plants were exposed to developmental temperatures atypical to the source environment.
• Seed and seedling traits suggest tetraploids are generally better adapted to stressful environments than diploids. Because tetraploid seeds of T. triandra are more dormant they are less likely to germinate under stress, and when they do germinate, seedling growth is rapid and independent of seed developmental environment. These novel results demonstrate that polyploidy, sometimes in interaction with developmental environment and possibly also asexuality, can have within‐species variation in seed and seedling traits that increase fitness in stressful environments.

     

Rainfall seasonality predicts the germination behavior of a tropical dry‐forest vine

Seed dormancy is considered to be an adaptive strategy in seasonal and/or unpredictable environments because it prevents germination during climatically favorable periods that are too short for seedling establishment. Tropical dry forests are seasonal environments where seed dormancy may play an important role in plant resilience and resistance to changing precipitation patterns. We studied the germination behavior of seeds from six populations of the Neotropical vine Dalechampia scandens (Euphorbiaceae) originating from environments of contrasting rainfall seasonality. Seeds produced by second greenhouse‐generation plants were measured and exposed to a favorable wet environment at different time intervals after capsule dehiscence and seed dispersal. We recorded the success and the timing of germination. All populations produced at least some dormant seeds, but seeds of populations originating from more seasonal environments required longer periods of after‐ripening before germinating. Within populations, larger seeds tended to require longer after‐ripening periods than did smaller seeds. These results indicate among‐population genetic differences in germination behavior and suggest that these populations are adapted to local environmental conditions. They also suggest that seed size may influence germination timing within populations. Ongoing changes in seasonality patterns in tropical dry forests may impose strong selection on these traits.     

Effects of primary seed dormancy on lifetime fitness of Arabidopsis thaliana in the field

Background and AimsSeed dormancy determines the environmental niche of plants in seasonal environments, and has consequences for plant performance that potentially go far beyond the seed and seedling stages. In this study, we examined the cascading effects of seed dormancy on the expression of subsequent life-history traits and fitness in the annual herb Arabidopsis thaliana.MethodsWe planted seeds of >200 recombinant inbred lines (RILs) derived from a cross between two locally adapted populations (Italy and Sweden), and both parental genotypes at the native site of the Swedish population in three consecutive years. We quantified the relationship between primary seed dormancy and the expression of subsequent life-history traits and fitness in the RIL population with path analysis. To examine the effects of differences in dormancy on the relative fitness of the two parental genotypes, we planted dormant seeds during the seed dispersal period and non-dormant seeds during the germination period of the local population.Key ResultsIn the RIL population, strong primary dormancy was associated with high seedling survival, but with low adult survival and fecundity, and path analysis indicated that this could be explained by effects on germination timing, rosette size and flowering start. The relationship between primary seed dormancy and germination proportion varied among years, and this was associated with differences in seasonal changes in soil moisture. The planting of dormant and non-dormant seeds indicated that the lower primary dormancy of the local Swedish genotype contributed to its higher germination proportion in two years and to its higher fecundity in one year.ConclusionsOur results show that seed dormancy affects trait expression and fitness components across the life cycle, and suggest that among-year variation in the incidence of drought during the germination period should be considered when predicting the consequences of climatic change for population growth and evolution.     

The role of domestication and maternal effects on seed traits of crop–wild sunflower hybrids (Helianthus annuus)

Hybridisation between crops and their wild relatives may promote the evolution of weeds. Seed germination and dormancy are the earliest life‐history traits and are highly influenced by the maternal parent. However, the ecological role of the maternal effect on seed traits in the evolution of crop–wild hybrids has received little attention. In this study, we test the relative importance of maternal and hybridisation effects on seed traits of the first generation of crop–wild sunflower hybrids (Helianthus annuus). Seed germination was tested in two wild populations with contrasting dormancy, two cultivated materials and their reciprocal crosses at four different times after harvest and three different temperatures. Seed germination at each of the four times, after ripening response and secondary dormancy were recorded along with four morphological traits. Additionally, the pericarp anatomy was analysed with light and scanning electron microscopy. We observed strong maternal effects on all seed traits. Seed germination, morphology and pericarp anatomy differed largely between the crop and wild seeds and these traits in the crop–wild hybrids resembled their female parent. Slight but significant hybridisation effects were observed in germination, mainly in seeds produced on wild plants. Crop hybridisation changed seed germination, the after ripening response and secondary dormancy in the crop direction. Morphological and anatomical traits associated with domestication strongly correlated with the observed differences in seed germination and dormancy in crop–wild sunflower hybrids. The large maternal effects along with the evolutionary divergence in seed traits were responsible for the large phenotypic differences observed in crop–wild hybrids with the same genetic composition. Wild‐like seed traits of hybrids suggest that there are no barriers to crop gene introgression at the seed level whereas crop‐like seed traits could be strongly selected against, conditioning the selection of traits expressed later in the life cycle and in the next generations.     

生长素调控种子的休眠与萌发

植物种子的休眠与萌发,是植物生长发育过程中的关键阶段,也是生命科学领域的研究热点。种子从休眠向萌发的转换是极为复杂的生物学过程,由外界环境因子、体内激素含量及信号传导和若干关键基因协同调控。大量研究表明,植物激素脱落酸(Abscisic acid, ABA)和赤霉素(Gibberellin acid, GA)是调控种子休眠水平,决定种子从休眠转向萌发的主要内源因子。ABA与GA在含量和信号传导两个层次上的精确平衡,确保了植物种子能以休眠状态在逆境中存活,并在适宜的时间启动萌发程序。生长素(Auxin)是经典植物激素之一,其对向性生长和组织分化等生物学过程的调控已有大量研究。但最近有研究证实,生长素对种子休眠有正向调控作用,这表明生长素是继ABA之后的第二个促进种子休眠的植物激素。本文在回顾生长素的发现历程、阐释生长素体内合成途径及信号传导通路的基础上,重点综述了生长素通过与ABA的协同作用调控种子休眠的分子机制,并对未来的研究热点进行了讨论和展望。     

Dormancy and germination: making every seed count in restoration

From 50 to 90% of wild plant species worldwide produce seeds that are dormant upon maturity, with specific dormancy traits driven by species' occurrence geography, growth form, and genetic factors. While dormancy is a beneficial adaptation for intact natural systems, it can limit plant recruitment in restoration scenarios because seeds may take several seasons to lose dormancy and consequently show low or erratic germination. During this time, seed predation, weed competition, soil erosion, and seed viability loss can lead to plant re‐establishment failure. Understanding and considering seed dormancy and germination traits in restoration planning are thus critical to ensuring effective seed management and seed use efficiency. There are five known dormancy classes (physiological, physical, combinational, morphological, and morphophysiological), each requiring specific cues to alleviate dormancy and enable germination. The dormancy status of a seed can be determined through a series of simple steps that account for initial seed quality and assess germination across a range of environmental conditions. In this article, we outline the steps of the dormancy classification process and the various corresponding methodologies for ex situ dormancy alleviation. We also highlight the importance of record‐keeping and reporting of seed accession information (e.g. geographic coordinates of the seed collection location, cleaning and quality information, storage conditions, and dormancy testing data) to ensure that these factors are adequately considered in restoration planning.     

Selection for low dormancy in annual ryegrass (Lolium rigidum) seeds results in high constitutive expression of a glucose-responsive α-amylase isoform

Background and Aims

α-Amylase in grass caryopses (seeds) is usually expressed upon commencement of germination and is rarely seen in dry, mature seeds. A heat-stable α-amylase activity was unexpectedly selected for expression in dry annual ryegrass (Lolium rigidum) seeds during targeted selection for low primary dormancy. The aim of this study was to characterize this constitutive activity biochemically and determine if its presence conferred insensitivity to the germination inhibitors abscisic acid and benzoxazolinone.

Methods

α-Amylase activity in developing, mature and germinating seeds from the selected (low-dormancy) and a field-collected (dormant) population was characterized by native activity PAGE. The response of seed germination and α-amylase activity to abscisic acid and benzoxazolinone was assessed. Using an alginate affinity matrix, α-amylase was purified from dry and germinating seeds for analysis of its enzymatic properties.

Key Results

The constitutive α-amylase activity appeared late during seed development and was mainly localized in the aleurone; in germinating seeds, this activity was responsive to both glucose and gibberellin. It migrated differently on native PAGE compared with the major activities in germinating seeds of the dormant population, but the enzymatic properties of α-amylase purified from the low-dormancy and dormant seeds were largely indistinguishable. Seed imbibition on benzoxazolinone had little effect on the low-dormancy seeds but greatly inhibited germination and α-amylase activity in the dormant population.

Conclusions

The constitutive α-amylase activity in annual ryegrass seeds selected for low dormancy is electrophoretically different from that in germinating seeds and its presence confers insensitivity to benzoxazolinone. The concurrent selection of low dormancy and constitutive α-amylase activity may help to enhance seedling establishment under competitive conditions.     

The role of invertebrates in seedling establishment in a heterocarpic plant,Atriplex sagittata

Seed heteromorphism is a marked character of many Chenopodioideae (Amaranthaceae). Seed morphs differ in dormancy, germination and seedling biology, but differences in their predation have not yet been studied. Atriplex sagittata produces small black dormant and large brown non‐dormant seeds. In this study, the timing of seed release and seedling establishment were ascertained, and their consumption by invertebrates (carabids, isopods and slugs) was studied. Seeds dispersed in the autumn passed the winter on the ground surface, protected from invertebrate predation by low temperatures. In the following vegetative season, ungerminated black seeds exposed to predation on ground surface were preferred by a large carabid species, Pseudoophonus rufipes. Some black and all brown seeds escaped predation by germinating in early spring. The seedlings were little endangered by carabids and isopods but were preferred by an invasive slug, Arion vulgaris, the feeding of which can exterminate seedlings at places in which slugs are abundant. Invertebrate predation is important factor of seed and seedling mortality of A. sagittata and seed heteromorphism modifies its intensity and timing.     

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

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

ENVIRONMENTAL AND GENETIC MATERNAL EFFECTS ON SEED CHARACTERS IN NEMOPHILA MENZIESII

Nuclear genetic, maternal genetic and maternal environmental effects on seed characters were estimated in the California native annual plant Nemophila menziesii using two greenhouse crosses. In one cross, according to a nested mating design, the narrow sense heritability of seed weight was small (3.9%). A subset of full-sib progenies produced in this cross was grown singly and in competition with the introduced grass Bromus diandrus. In a second cross, these plants were used as mothers (dams) and were each mated to the same three sires. Seeds produced by mothers competing with B. diandrus showed a significant reduction in weight, increase in time to germination, and increase in the incidence of dormancy, when compared to seeds from mothers grown singly. Significant sire components were found for time to germination and incidence of dormancy. Maternal genetic variation for seed weight was largely expressed as maternal genotype by maternal environment interaction, and showed no significant maternal genetic main effect. Time to germination and dormant fraction showed a relatively large maternal genetic effect. Evolution of seed characters in N. menziesii is more likely to occur via indirect response to selection among maternal plants than among the seeds themselves. Maternal genotype by maternal environment interaction could potentially contribute to the maintenance of maternal genetic variation in seed weight, but this does not appear likely for dormancy.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.     

皇后葵及香棕种子休眠机理与催芽技术探讨

本文以生产上发芽不整齐的皇后葵和香棕种子为材料,研究棕榈科植物种子的休眠机理及催芽技术。试验结果表明,皇后葵及香棕的果肉及胚乳中均含有发芽抑制物质,同一植株的种子发芽率及发芽力的差别还与种子的成熟度有关,用GA₃100mg/L或NAA 100mg/L浸种有利于提高发芽率。     

PHENOTYPIC CONSEQUENCES OF FORCING GERMINATION: A GENERAL PROBLEM OF INTERVENTION IN EXPERIMENTAL DESIGN

Many evolutionary and ecological studies of plants with seed dormancy are plagued by design problems related to seed germination. On the one hand, traits of interest, especially life history traits like early growth rate or time to flowering, may not be independent of dormancy phenotype. On the other hand, germination-inducing treatments are likely to affect these traits of interest. Consequently, researchers often have had to use confounded designs without information about the consequences. To examine this problem, we studied how early growth in California poppy (Eschscholzia californica) is affected by treating seeds with giberellic acid (GA) to stimulate germination. The dose used was sufficient to slightly but significantly advance emergence time in treated seeds, but it appeared to cause variable growth responses among treated seeds. Experiments using GA to stimulate germination may thus be misleading, at least for the populations and dose studied. Perhaps more importantly, the experimental and statistical approach we used can be employed to study the effects of this confounded design for other doses and 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.     

Seed development and germination ecophysiology of the invasive tree <Emphasis Type="Italic">Prunus serotina</Emphasis> (Rosaceae) in a temperate forest in Western Europe

Seed development, dormancy and germination of the American invasive tree species, Prunus serotina, are described for plants growing in a large forest in Belgium. Seeds of P. serotina were collected following anthesis in the first week of July and thereafter at fortnightly intervals. Seed dormancy, temperature requirements for germination and the soil seed bank were investigated. At maturation (about 105 days after anthesis), seed moisture content had decreased to around 13.7%, and 44% of the seeds had attained the capacity to germinate. Mature seeds of P. serotina exhibited physiological dormancy, germinating only after a long cold, moist stratification period. Highest germination percentage occurred in seeds treated with gibberellic acid (GA₃), at 10°C. We found no evidence that P. serotina forms a persistent seed bank but noticed a persistent seedling bank in the field.     

西南獐芽菜种子萌发对温度、光照和外源生长调节物质的响应

研究不同温度、光照和外源生长调节物质处理种子对西南獐芽菜种子萌发特性的结果表明,西南獐芽菜种子为休眠型种子,其最适合萌发温度为20℃。光照促进种子的萌发,但影响并不显著。低浓度（10mg·L-1）6-BA促进西南獐芽菜种子萌发,而高浓度（50mg·L-1）的则抑制其萌发,不同浓度和浸种时间的赤霉素（GA）和IAA均促进种子萌发,GA的效果更好。     

Ethylene as a Component of the Emanations From Germinating Peanut Seeds and Its Effect on Dormant Virginia-type Seeds

The embryonic axes of Spanish-type peanut seeds that do not exhibit dormancy to any extent were found to produce ethylene during germination. Virginia-type peanut seeds of the extremely dormant variety NC-13 produced low levels of ethylene when imbibed but not germinating. Treatments that released dormancy of NC-13 peanut seeds resulted in increased ethylene production by the embryonic axis. The estimated internal concentration of ethylene in Virginia-type peanut seeds was 0.4 ppm at 24 hr of germination. Fumigation with an external concentration of 3.0 to 3.5 ppm for 6 hr was sufficient to break dormancy of Virginia-type peanut seeds. These results suggest that ethylene is associated with the germination processes of non-dormant seeds and participates in the breaking of seed dormancy of dormant peanut varieties.     

AtPER1 enhances primary seed dormancy and reduces seed germination by suppressing the ABA catabolism and GA biosynthesis in Arabidopsis seeds

Seed is vital to the conservation of germplasm and plant biodiversity. Seed dormancy is an adaptive trait in numerous seed‐plant species, enabling plants to survive under stressful conditions. Seed dormancy is mainly controlled by abscisic acid (ABA) and gibberellin (GA) and can be classified as primary and secondary seed dormancy. The primary seed dormancy is induced by maternal ABA. Here we found that AtPER1, a seed‐specific peroxiredoxin, is involved in enhancing primary seed dormancy. Two loss‐of‐function atper1 mutants, atper1‐1 and atper1‐2, displayed suppressed primary seed dormancy accompanied with reduced ABA and increased GA contents in seeds. Furthermore, atper1 mutant seeds were insensitive to abiotic stresses during seed germination. The expression of several ABA catabolism genes (CYP707A1, CYP707A2, and CYP707A3) and GA biosynthesis genes (GA20ox1, GA20ox3, and KAO3) in atper1 mutant seeds was increased compared to wild‐type seeds. The suppressed primary seed dormancy of atper1‐1 was completely reduced by deletion of CYP707A genes. Furthermore, loss‐of‐function of AtPER1 cannot enhance the seed germination ratio of aba2‐1 or ga1‐t, suggesting that AtPER1‐enhanced primary seed dormancy is dependent on ABA and GA. Additionally, the level of reactive oxygen species (ROS) in atper1 mutant seeds was significantly higher than that in wild‐type seeds. Taken together, our results demonstrate that AtPER1 eliminates ROS to suppress ABA catabolism and GA biosynthesis, and thus improves the primary seed dormancy and make the seeds less sensitive to adverse environmental conditions.