Interaction of maternal environment and allelic differences in seed vigour genes determines seed performance in Brassica oleracea

Seed vigour is a key trait essential for the production of sustainable and profitable crops. The genetic basis of variation in seed vigour has recently been determined in Brassica oleracea, but the relative importance of the interaction with parental environment is unknown. We produced seeds under a range of maternal environments, including global warming scenarios. Lines were compared that had the same genetic background, but different alleles (for high and low vigour) at the quantitative trait loci responsible for determining seed vigour by altering abscisic acid (ABA) content and sensitivity. We found a consistent effect of beneficial alleles across production environments; however, environmental stress during production also had a large impact that enhanced the genetic difference in seed performance, measured as germination speed, resistance to controlled deterioration and induction of secondary dormancy. Environmental interaction with allelic differences in key genes that determine ABA content and sensitivity develops a continuity in performance from rapid germination through to failure to complete germination, and increasing depths of seed dormancy. The genetic–environmental interaction revealed provides a robust mechanism of bet‐hedging to minimize environmental risk during subsequent germination, and this could have facilitated the rapid change in seed behaviour (reduced dormancy and rapid germination) observed during crop domestication.     

Morphological and functional seed traits of the wild medicinal plant Dioscorea strydomiana,the most threatened yam in the world

• Morphological and functional seed traits have important roles in characterising the species regeneration niche and help to understand the reproductive biology of rare and threatened plants, which can thus support appropriate plant conservation measures.
• Seed morphometric and dispersal kinetics of the critically endangered Dioscorea strydomiana were measured and compared with those of four other Dioscorea species, and seed germination response under constant temperatures (5–35 °C) was compared with that of the congeneric and widespread D. sylvatica.
• Seed mass of D. strydomiana (ca. 14 mg) was twice that of D. sylvatica, but similar to or smaller than the other species examined. Seeds of D. strydomiana have the lowest speed of descent and lowest variability in most of the morphological traits considered, suggesting lower phenotypic plasticity but higher variance in the wing‐loading value. Seeds of D. strydomiana reached maximum germination at 15 °C (ca. 47%), which decreased slightly to ca. 37% at 25 °C and was completely inhibited at 35 °C. D. sylvatica seeds started to germinate at 10 °C (ca. 3%), reached 75–80% germination at 15–20 °C and maximum (ca. 90%) at 25–30 °C. Base temperatures for germination (T_b) were 9.3 and 5.7 °C, for D. strydomiana and D. sylvatica, respectively. Due to the higher germination percentages of D. sylvatica, ceiling and optimum temperatures could also be modelled for this species, suggesting higher sensitivity to high temperature for seeds of D. strydomiana.
• The detected poor seed lot quality of D. strydomiana suggests difficulties in reproduction from seed, highlighting the need for further investigation and conservation actions for this threatened yam species.

     

Optimising seed processing techniques to improve germination and sowability of native grasses for ecological restoration

• Grasslands across the globe are undergoing expansive degradation due to human impacts and climate change. If restoration of degraded native grassland is to be achieved at the scale now required, cost‐effective means for seed‐based establishment of grass species is crucial. However, grass seeds present numerous challenges associated with handling and germination performance that must be overcome to improve the efficiency of seeding. Previous research has demonstrated that complete removal of the palea and lemma (husk) maximises germination performance, hence we investigated the effects of complete husk removal on seed handling and germination of four temperate Australian grass species.
• Three techniques were tested to remove the husk – manual cleaning, flaming or acid digestion (the latter two followed by a manual cleaning step); these techniques were refined and adapted to the selected species, and germination responses were compared.
• The complete removal of the husk improved seed handling and sowability for all species. Germination was improved in Microlaena stipoides by 19% and in Rytidosperma geniculatum by 11%. Of the husk removal methods tested, flaming was detrimental to seed germination and fatal for one species (R. geniculatum). Compared to manual cleaning, sulphuric acid improved the overall efficacy of the cleaning procedure and increased germination speed (T50) in Austrostipa scabra, Chloris truncata and M. stipoides, and improved final germination in R. geniculatum by 13%.
• The seed processing methods developed and tested in the present study can be applied to grass species that present similar handling and germination performance impediments. These and other technological developments (seed coating and precision sowing) will facilitate more efficient grassland restoration at large scale.

     

Comparative germination responses to water potential across different populations of Aegilops geniculata and cultivar varieties of Triticum durum and Triticum aestivum

• Crop Wild Relatives are often used to improve crop quality and yields because they contain genetically important traits that can contribute to stress resistance and adaptation.
• Seed germination of different populations of Aegilops geniculata Roth collected along a latitudinal gradient was studied under different drought stress in order to find populations suitable for improving drought tolerance in wheat. Different accessions of Aegilops neglecta Req. ex Bertol., Triticum aestivum L. and T. durum Desf. were used as comparison.
• Under full hydration, germination was high in all populations, but increasing drought stress led to reduced and delayed germination. Significant differences in final germination and mean time to germinate were detected among populations. Wheat, durum wheat and the southern population of Ae. geniculata were not significantly affected by drought stress, germinating similarly under all treatments. However, seed germination of the northern populations of Ae. geniculata was significantly reduced under high water stress treatment. Differences between populations of the same species could not be explained by annual rainfall across populations’ distributions, but by rainfall during seed development and maturation.
• Differences in the germination responses to drought found here highlight the importance of source populations as criteria for genotype selection for pre‐breeders.

     

Comparison of seed and seedling functional traits in native Helianthus species and the crop H. annuus (sunflower)

• Seed functional traits of native Helianthus species contribute towards ecosystem services but limitations to their use in managed programmes exist. Many perennial Helianthus possess seed dormancy. The ability for germination to occur under different temperature and drought conditions, as well as the capacity of germinated seeds to convert into normal seedlings is rarely considered. Our aim was to identify and quantify these constraints through functional trait analyses.
• In five seed lots of native Helianthus (four perennial and one annual) and five genotypes of sunflower (H. annuus) for comparison, dormancy, thermal and hydro thresholds and times, morphology, mass, oil content and conversion into normal seedlings were quantified. The influence of the seed collection site environment on these traits was also explored.
• Seed dormancy of the perennial species was overcome by scarification followed by germination in 5 mm GA₃. Thermal and hydro‐time analyses revealed slower germination for the native seed lots (>1350 °Ch) in comparison to the sunflower genotypes (<829.9 °Ch). However, native seed lots had a higher capacity to convert into normal seedlings at high temperatures and low water potentials than sunflower genotypes. For the native seed lots, the average monthly temperature of the collection site was negatively correlated with thermal time.
• Variability in seed functional traits of native Helianthus and greater capacity for germinated seeds to convert into normal seedlings suggests they are better equipped to cope with high temperature and drought scenarios than sunflower. Effective dormancy alleviation is required to facilitate the use of native Helianthus species.

     

Effect of maturation conditions on light and temperature requirements during seed germination of Citrullus colocynthis from the Arabian Desert

• Seed germination of Citrullus colocynthis, as in many other species of Cucurbitaceae, is inhibited by light, particularly at low temperatures. Germination response to light and temperature has been attributed to day length and temperature during seed maturation. This study assessed the effects of these factors on the germination response of C. colocynthis to temperature and light quality.
• Ripe fruits were collected from natural habitats during December and February and germinated at three temperatures (15/25, 20/30 and 25/35 °C) in five light treatments (dark, white light and Red:Far Red (R:FR) ratios of 0.30, 0.87 and 1.19). Additionally, unripe fruits were also collected from natural habitats and completed their maturation in growth chambers under different day lengths (6, 16 and 24 h of darkness) at 10/20 °C, and in darkness at both 10/20 °C and 25/35 °C. Mature seeds of the different treatments were germinated in the same five light treatments at 15/25 °C.
• Germination was significantly higher in the dark than that in any light treatment. Seeds matured at higher temperatures (i.e. seeds from the December collection and those matured at 25/35 °C) had significantly higher germination than those matured at lower temperatures (i.e. seeds from the February collection and those matured at 10/20 °C). Dark germination was significantly higher for the December collection than for the February collection. Seeds of the two collections germinated in the dark only at 15/25 °C. However, seeds matured in a growth chamber at 10/20 °C in darkness germinated at 15/25 °C in all light treatments, except for the R:FR ratio 0.30. Seeds of the different treatments failed to germinate in FR‐rich light.
• This study demonstrates that both temperature and day length during seed maturation play significant roles in the germination response of C. colocynthis. Additionally, the dark requirement for germination is likely beneficial for species with the larger seeds, such as C. colocynthis, which produce bigger seedlings that are able to emerge from deep soils and are competitively superior under dense vegetation and resource‐limited conditions.

     

Heteromorphic seeds of wheat wild relatives show germination niche differentiation

• Crop wild relatives are fundamental genetic resources for crop improvement. Wheat wild relatives often produce heteromorphic seeds that differ in morphological and physiological traits. Several Aegilops and Triticum species possess, within the same spikelet, a dimorphic seed pair, with one seed being larger than the other. A comprehensive analysis is needed to understand which traits are involved in seed dimorphism and if these aspects of variation in dimorphic pairs are functionally related.
• To this end, dispersal units of Triticum urartu and five Aegilops species were X‐rayed and the different seed morphs weighed. Germination tests were carried out on seeds, both dehulled and left in their dispersal units. Controlled ageing tests were performed to detect differences in seed longevity among seed morphs, and the antioxidant profile was assessed in terms of antioxidant compounds equipment and expression of selected antioxidant genes. We used PCA to group seed morphs sharing similar patterns of germination traits, longevity estimates and antioxidant profile.
• Different seed morphs differed significantly in terms of mass, final germination, germination timing, longevity estimates and antioxidant profile in most of the tested species. Small seeds germinated slower, had lower germination when left in their dispersal units, a higher antioxidant potential and were longer‐lived than large seeds. The antioxidant gene expression varied between morphs, with different patterns across species but not clearly reflecting the phenotypic observations.
• The results highlight different trait trade‐offs in dimorphic seeds of Aegilops and T. urartu, affecting their germination phenology and longevity, thereby resulting in recruitment niche differentiation.

     

Estimating seed dispersal distance: A comparison of methods using animal movement and plant genetic data on two primate‐dispersed Neotropical plant species

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Protocols for Use of Potamogeton perfoliatus and Ruppia maritima Seeds in Large‐Scale Restoration

Restoration of submerged aquatic vegetation from seed has been hampered by a lack of information on the appropriate conditions for collecting, processing, and storing seeds prior to dispersal. Seeds must be processed and stored under conditions that maintain seed viability, meet dormancy requirements, and prevent premature germination. This study examined the effects of collection date, processing technique, aeration, storage and induction temperature and salinity, and storage period on seed germination of two mesohaline aquatic species, Potamogeton perfoliatus and Ruppia maritima. Collection date and processing technique were significant factors affecting seed yield from donor populations. Seeds of both species remained viable and germinated best when stored at 4°C, and then exposed to freshwater induction conditions. However, their responses to other factors differed. Aeration during storage was necessary in order to maintain viability of P. perfoliatus seeds, whereas it was unnecessary for R. maritima seeds. Storage in freshwater at 4°C prevented germination of P. perfoliatus seeds, while high salinity during cold storage was necessary to minimize premature germination of R. maritima. Mean germination time of P. perfoliatus was dependent on storage salinity; in contrast, mean germination time of R. maritima seeds was dependent on induction salinity. These differences indicate that the methods required to produce large quantities of underwater plant seed amenable to large‐scale restoration efforts must be tailored to the specific requirements of individual species and must consider the range of processes from initial harvest through seed testing prior to field establishment.     

Ecological longevity of Polaskia chende (Cactaceae) seeds in the soil seed bank,seedling emergence and survival

• Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5‐year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico.
• P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12‐h photoperiod under white light, far‐red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years.
• The species forms long‐term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling.
• Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence.

     

Seed dormancy and control of germination in Sisymbrella dentata (L.) O.E. Schulz (Brassicaceae)

• Seed germination responsiveness to environmental cues is crucial for plant species living in changeable habitats and can vary among populations within the same species as a result of adaptation or modulation to local climates. Here, we investigate the germination response to environmental cues of Sisymbrella dentata (L.) O.E. Schulz, an annual endemic to Sicily living in Mediterranean Temporary Ponds (MTP), a vulnerable ecosystem.
• Germination of the only two known populations, Gurrida and Pantano, was assessed over a broad range of conditions to understand the role of temperatures, nitrate, hormones (abscisic acid – ABA and gibberellins – GA) and after‐ripening in dormancy release in this species.
• Seed germination responsiveness varied between the two populations, with seeds from Gurrida germinating under a narrower range of conditions. Overall, this process in S. dentata consisted of testa and endosperm rupture as two sequential events, influenced by ABA and GA biosynthesis. Nitrate addition caused an earlier testa rupture, after‐ripening broadened the thermal conditions that allow germination, and alternating temperatures significantly promoted germination of non‐after‐ripened seeds.
• Primary dormancy in S. dentata seeds likely allows this plant to form a persistent seed bank that is responsive to specific environmental cues characteristic of MTP habitats.

     

Seed germination and dormancy traits of forbs and shrubs important for restoration of North American dryland ecosystems

• In degraded dryland systems, native plant community re‐establishment following disturbance is almost exclusively carried out using seeds, but these efforts commonly fail. Much of this failure can be attributed to the limited understanding of seed dormancy and germination traits.
• We undertook a systematic classification of seed dormancy of 26 species of annual and perennial forbs and shrubs that represent key, dominant genera used in restoration of the Great Basin ecosystem in the western United States. We examined germination across a wide thermal profile to depict species‐specific characteristics and assessed the potential of gibberellic acid (GA₃) and karrikinolide (KAR₁) to expand the thermal germination envelope of fresh seeds.
• Of the tested species, 81% produce seeds that are dormant at maturity. The largest proportion (62%) exhibited physiological (PD), followed by physical (PY, 8%), combinational (PY + PD, 8%) and morphophysiological (MPD, 4%) dormancy classes. The effects of chemical stimulants were temperature‐ and species‐mediated. In general, mean germination across the thermal profile was improved by GA₃ and KAR₁ for 11 and five species, respectively. We detected a strong germination response to temperature in freshly collected seeds of 20 species. Temperatures below 10 °C limited the germination of all except Agoseris heterophylla, suggesting that in their dormant state, the majority of these species are thermally restricted.
• Our findings demonstrate the utility of dormancy classification as a foundation for understanding the critical regenerative traits in these ecologically important species and highlight its importance in restoration planning.

     

Seed dispersal in the mycoheterotrophic orchid Yoania japonica: Further evidence for endozoochory by camel crickets

• Although orthopterans are rarely considered to be effective seed dispersal agents, the large flightless crickets known as ‘weta’ have been suggested to function as ecological replacements for small mammals in New Zealand, where such mammals are absent. In addition, a recent study reported that camel crickets mediate seed dispersal of several heterotrophic plants, including Yoania amagiensis in Japan.
• I investigated the seed dispersal mechanism of Yoania japonica because the fruit morphology is similar to Y. amagiensis. Specifically, I aimed to determine whether Y. japonica fruits are consumed by camel crickets and, if so, whether the seeds defecated by camel crickets remains intact, by checking seed viability with TTC staining, and whether germination rate is different between seeds collected directly from fruits and defecated seeds by comparing in situ seed germinability.
• The present study provides evidence that camel crickets function as seed dispersal agents of Y. japonica. Camel crickets were important consumers of Y. japonica fruits, and a substantial portion of the consumed seeds remained viable after passing through the digestive tract. In situ seed germination experiments revealed that the seeds defecated by camel crickets actually germinated in the field. In addition, the germination rate of defecated seeds was even higher than that of intact seeds, although the difference was not significant.
• Taken together with recent reports of insect‐mediated endozoochory, such a seed dispersal system may be common in plants with fleshy indehiscent fruits and small seeds, even in locations where other seed dispersal agents are present.

     

Molecular markers provide evidence for a broad‐fronted recolonisation of the widespread calcareous grassland species Sanguisorba minor from southern and cryptic northern refugia

• Calcareous grasslands belong to the most species‐rich and endangered habitats in Europe. However, little is known about the origin of the species typically occurring in these grasslands. In this study we analysed the glacial and post‐glacial history of Sanguisorba minor, a typical plant species frequently occurring in calcareous grasslands.
• The study comprised 38 populations throughout the whole distribution range of the species across Europe. We used molecular markers (AFLP) and applied Bayesian cluster analysis as well as spatial principal components analysis (sPCA) to identify glacial refugia and post‐glacial migration routes to Central Europe.
• Our study revealed significant differences in the level of genetic variation and the occurrence of rare fragments within populations of S. minor and a distinct separation of eastern and western lineages. The analyses uncovered traditional southern but also cryptic northern refugia and point towards a broad fronted post‐glacial recolonisation.
• Based on these results we postulate that incomplete lineage sorting may have contributed to the detected pattern of genetic variation and that S. minor recolonised Central Europe post‐glacially from Iberia and northern glacial refugia in France, Belgium or Germany. Our results highlight the importance of refugial areas for the conservation of intraspecific variation in calcareous grassland species.

     

Quantitative and qualitative consequences of reduced pollen loads in a mixed‐mating plant

Greater pollination intensity can enhance maternal plant fitness by increasing seed set and seed quality as a result of more intense pollen competition or enhanced genetic sampling. We tested experimentally these effects by varying the pollen load from a single pollen donor on stigmas of female flowers of Dalechampia scandens (Euphorbiaceae) and measuring the effects on seed number and seed mass. Seed set increased rapidly with pollen number at low to moderate pollen loads, and a maximum set of three seeds occurred with a mean pollen load of 19 pollen grains. We did not detect a trade‐off between the number of seeds and seed mass within a fruit. Seed mass increased with increasing pollen load, supporting the hypothesis of enhanced seed quality via increased pollen‐competition intensity or genetic sampling. These results suggest that maternal fitness increases with larger pollen loads, even when the fertilization success is already high. Our results further highlight the importance of high rates of pollen arrival onto stigmas, as mediated by reliable pollinators. Comparing the pollen‐to‐seed response curve obtained in this experiment with those observed in natural populations suggests that pollen limitation may be more severe in natural populations than predicted from greenhouse studies. These results also indicate that declines in pollinator abundance may decrease plant fitness through lowered seed quality before an effect on seed set is detected.     

Sowing seasons and drying methods during post-harvest influence the seed vigour of soybean (Glycine max (L.) Merr.)

Experiments were conducted to study the influence of sowing seasons and drying methods on the seed vigour of two spring soybean (Glycine max (L.) Merr.) cultivars. Two cultivars, ‘Huachun18’ and ‘Huachun 14’, were sown in three seasons viz., spring, summer and autumn and the harvested seeds were dried using three different methods. The results showed that soybean sown in spring had a higher number of branches per plant, pods per branch and seed weight, and consequently resulted in higher seed yields than that of soybean sown in autumn or summer seasons. Seeds sown in the autumn season had the lowest values of electrical conductivity during seed imbibitions, higher peroxidase (POD) activity in germinated seedlings and lower contamination by the seed-borne fungi on the MS medium, which indirectly improved the seed vigour, which was followed by summer sown seeds. Seeds sown during the spring season resulted in poor seed vigour. In addition, the effect of drying methods on the seed vigour was also clarified. Seeds that hung for four days before threshing and then air-dried had the poorest seed vigour which was determined by germination, electrical conductivity, POD activity and seed borne fungal growth. There was no difference in seed vigour between other methods, i.e. seeds threshed directly at harvest and then air-dried on a bamboo sifter or concrete floor. These results indicated that autumn sowing soybean and the drying method in which seeds were threshed directly at harvest and then air-dried on a bamboo sifter resulted in higher seed vigour.     

Efficient Detection of Leptosphaeria maculans from Infected Seed lots of Oilseed Rape

An efficient DNA extraction protocol and polymerase chain reaction (PCR) assay for detecting Leptosphaeria maculans from infected seed lots of oilseed rape were developed. L. maculans, the causal agent of blackleg, a damaging disease in oilseeds rape/canola worldwide, was listed as a quarantine disease by China in 2009. China imports several millions of tons of oilseeds every year. So there is a high risk that this pathogen will be introduced to China via contaminated seeds. Seed contamination is one of the most significant factors in the global spread of phytopathogens. Detection of L. maculans in infected seed lots by PCR assay is difficult due to the low level of pathogen mycelium/spores on seeds and PCR inhibitors associated with the seeds of oilseed rape. In our study, these two major obstacles were overcome by the development of a two‐step extraction protocol combined with a nested PCR. This extraction protocol (kit extraction after CTAB method) can efficiently extract high‐quality DNA for PCR. Amplification results showed that the detection threshold for conventional PCR and nested PCR was, respectively, 1 ng and 10 fg of DNA per μl in mycelia samples. On contaminated seed lots of oilseed rape, the detection threshold of conventional and nested PCR was 709 fg/μl and 709 ag/μl of DNA, respectively. The DNA extraction protocol and PCR assay developed in this study can be used for rapid and reliable detection of L. maculans from infected seeds of oilseed rape .     

Seed traits and germination behaviour of four Sardinian populations of Helichrysum microphyllum subsp. tyrrhenicum (Asteraceae) along an altitudinal gradient

• Helichrysum microphyllum subsp. tyrrhenicum (Asteraceae) is an endemic taxon of Sardinia and Corsica, where it grows at different altitudes. The objective of this study was to investigate the seed traits and germination behaviour of four Sardinian populations of this taxon located at different altitudes.
• Seed traits were evaluated, and germination tests were carried out by incubating seeds at a range of constant (5–30 °C) and alternating (25/10 °C) temperatures. The dry after‐ripening (DAR) pre‐treatment was also applied by storing seed in dry conditions for 3 months at 25 °C. Seed traits and germination behaviour data were statistically analysed to identify if there was a correlation with altitude.
• Differences in seed size, area and mass among populations were recorded, however, no relationship was found with altitude. High germination percentages were obtained in all populations, both in untreated and DAR seeds, and were positively affected by alternating temperatures. The final germination percentage and time required to reach 50% final germination (T₅₀) showed no relationship with altitude.
• The differences in seed traits and germination detected among the studied populations of H. microphyllum subsp. tyrrhenicum were not correlated with altitude. This study provides new and important knowledge for this taxon. H. microphyllum subsp. tyrrhenicum is characterised by high germination percentages and low T₅₀ values and does not seem to require any dormancy‐breaking treatment. This species represents a high‐potential native plant species that should be considered within environmental management plans.