Soil salinity regulates spatial-temporal heterogeneity of seed germination and seedbank persistence of an annual diaspore-trimorphic halophyte in northern China.

BACKGROUND AND AIMS: Seed heteromorphism is a plant strategy that an individual plant produces two or more distinct types of diaspores, which have diverse morphology, dispersal ability, ecological functions and different effects on plant life history traits. The aim of this study was to test the effects of seasonal soil salinity and burial depth on the dynamics of dormancy/germination and persistence/depletion of buried trimorphic diaspores of a desert annual halophyte Atriplex centralasiatica. METHODS: We investigated the effects of salinity and seasonal fluctuations of temperature on germination, recovery of germination and mortality of types A, B, C diaspores of A. centralasiatica in the laboratory and buried diaspores in situ at four soil salinities and three depths. Diaspores were collected monthly from the seedbank from December 2016 to November 2018, and the number of viable diaspores remaining (not depleted) and their germinability were determined. RESULTS: Non-dormant type A diaspores were depleted in the low salinity "window" in the first year. Dormant diaspore types B and C germinated to high percentages at 0.3 and 0.1 mol L-1 soil salinity, respectively. High salinity and shallow burial delayed depletion of diaspore types B and C. High salinity delayed depletion time of the three diaspore types and delayed dormancy release of types B and C diaspores from autumn to spring. Soil salinity modified the response of diaspores in the seedbank by delaying seed dormancy release in autum and winter and by providing a low-salt concentration window for germination of non-dormant diaspores in spring and early summer. CONCLUSIONS: Buried trimorphic diaspores of annual desert halophyte A. centralasiatica exhibited diverse dormancy/germination behavior in respond to seasonal soil salinity fluctuation. Prolonging persistence of the seedbank and delaying depletion of diaspores under salt stress in situ primarily is due to inhibition of dormancy-break. The differences in dormancy/germination and seed persistence in the soil seedbank may be a bet-hadging strategy adapted to stressful temporal and spatial heterogeneity, and allows A. centralasiatica to persist in the unpredictable cold desert enevironment.

Evolutionary reversal of physical dormancy to nondormancy: evidence from comparative seed morphoanatomy of Argyreia species (Convolvulaceae).

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

Rodents mediate the relationship between seed rain, seed bank, and plant community with increased grazing disturbance.

Seed rain and the soil seed bank represent the dispersal of seeds in space and time, respectively, and can be important sources of recruitment of new individuals during plant community regeneration. However, the temporal dynamics of seed rain and the mechanisms by which the seed rain and soil seed bank may play a role in plant community regeneration with increased grazing disturbance remain unclear. Seed rain, soil seed bank, aboveground vegetation, and rodent density were sampled along a grazing gradient in an alpine marsh on the eastern Tibetan Plateau. We described the temporal dynamics of seed dispersal using Bayesian generalized mixed models, and nonmetric multidimensional scaling and the structural equation model were used to examine the effects of grazing disturbance on the relative role of seed rain and soil seed bank on aboveground plant community regeneration. The temporal dynamics of seed rain changed from a unimodal to a bimodal pattern with increased grazing disturbance. Both species diversity and seed density of the seed rain and seed bank increased significantly with increased grazing disturbance. Increased grazing disturbance indirectly increased the similarity of composition between seed rain, seed bank, and aboveground plant community by directly increasing species diversity and abundance of aboveground plant community. However, increased grazing disturbance also indirectly decreased the similarity of seed rain, soil seed bank, and aboveground plant community by directly increasing rodent density. The similarity between seed rain and aboveground plant community was greater than that of the soil seed bank and aboveground plant community with increased grazing disturbance. Grazing disturbance spreads the risk of seed germination and seedling establishment by changing the temporal dynamics of seed dispersal. Plants (positive) and rodents (negative) mediated the role of seed rain and soil seed bank in plant community regeneration. The role of seed rain in plant community regeneration is higher than the seed bank in disturbed alpine marshes. Our findings increase our understanding of the regeneration process of the plant community, and they provide valuable information for the conservation and restoration of alpine marsh ecosystems.

Environmental and seed-position effects on viability and germination of buried seeds of an invasive diaspore-heteromorphic annual grass.

Environmental factors, such as temperature and moisture, and plant factors, such as seed position on the mother plant, can affect seed viability and germination. However, little is known about the viability and germination of seeds in different positions on the mother plant after burial in soil under natural environmental conditions. Here, diaspores from three positions on a compound spike and seeds from two/three positions in a diaspore of the invasive diaspore-heteromorphic annual grass Aegilops tauschii were buried at four depths for more than 2 years (1-26 months) under natural conditions and viability and germination monitored monthly. Viability of seeds in each diaspore/seed position decreased as burial depth and duration increased and was associated with changes in soil temperature and moisture. Germination was highest at 2 cm and lowest at 10 cm soil depths, with peaks and valleys in autumn/spring and winter/summer, respectively. Overall, seeds in distal diaspore and distal seed positions had higher germination percentages than those in basal diaspore and basal seed positions, but basal ones lived longer than distal ones. Chemical content of fresh diaspores/seeds was related to diaspore/seed position effects on seed germination and viability during burial. We conclude that seeds in distal diaspores/seed positions have a 'high risk' strategy and those in basal positions a 'low risk' strategy. The two risk strategies may act as a bet-hedging strategy that spreads risks of germination failure in the soil seed bank over time, thereby facilitating the survival and invasiveness of A. tauschii.

A long-term experiment reveals no trade-off between seed persistence and seedling emergence.

Information on seed persistence and seedling emergence from the soil seed bank is critical for understanding species coexistence and predicting community dynamics. However, quantifying seed persistence in the soil is challenging; thus, its association with other life-history traits is poorly known on a broad scale. Using germination phenology for 349 species in a 42-yr experiment, we quantified the persistence-emergence correlations and their associations with intrinsic regeneration traits using Bayesian phylogenetic multilevel models. We showed no trade-off between seed persistence and seedling emergence. Physically dormant seeds were more persistent but exhibited lower emergence than nondormant seeds. Monocarpic species had both higher persistence and emergence than polycarpic species. Seed mass posed a marginal proxy for persistence, while emergence almost doubled from the smallest to the largest seeds. This study challenges the traditional assumption and is the first demonstration of noncorrelation between persistence and emergence, probably owing to the complexity of regenerative strategies. Species with short persistence and low emergence would be the most vulnerable for in situ conservation. Our analyses of this unique, long-term dataset provide a strong incentive for further experimental studies and a rich data resource for future syntheses.

Inflated Ovary May Increase the Dispersal Ability of Three Species in the Cold Deserts of Central Asia.

Among the diaspores of angiosperms an inflated ovary (IO) is a novel morphological trait, but no studies have evaluated its effects on dispersal. The primary aim of this study was to determine the effect of the IO on diaspore dispersal in three cold desert species (Carex physodes, Calligonum junceum, and Sphaerophysa salsula). Various morphological features and the mass of fruits and seeds of each species were measured. The role of an IO in diaspore dispersal by wind and water was determined by comparing responses of intact (inflated) IOs and flattened fruits and seeds. Mature diaspores of three species were dispersed by wind, and the IO significantly increased dispersal distance in the field and at different wind speeds in the laboratory. The floating time on water was greater for inflated fruits than flattened fruits and seeds. Since the seed remains inside the IO until after dispersal is completed, the IO of the three species enhances diaspore dispersal. This is the first detailed study on how an IO increases diaspore/seed dispersal. Furthermore, after primary dispersal by wind, secondary dispersal can occur via wind or surface runoff of water, and each method is enhanced by the presence of an IO.

Global seed dormancy patterns are driven by macroclimate but not fire regime.

Seed dormancy maximizes plant recruitment in habitats with variation in environmental suitability for seedling establishment. Yet, we still lack a comprehensive synthesis of the macroecological drivers of nondormancy and the different classes of seed dormancy: physiological dormancy, morphophysiological dormancy and physical dormancy. We examined current geographic patterns and environmental correlates of global seed dormancy variation. Combining the most updated data set on seed dormancy classes for > 10 000 species with > 4 million georeferenced species occurrences covering all of the world's biomes, we test how this distribution is driven by climate and fire regime. Seed dormancy is prevalent in seasonally cold and dry climates. Physiological dormancy occurs in relatively dry climates with high temperature seasonality (e.g. temperate grasslands). Morphophysiological dormancy is more common in forest-dominated, cold biomes with comparatively high and evenly distributed precipitation. Physical dormancy is associated with dry climates with strong seasonal temperature and precipitation fluctuations (e.g. deserts and savannas). Nondormancy is associated with stable, warm and wetter climates (e.g. tropical rain forest). Pyroclimate had no significant effect on the distribution of seed dormancy. The environmental drivers considered in this study had a comparatively low predictive power, suggesting that macroclimate is just one of several global drivers of seed dormancy.

Variation in Seed Dormancy of Chaco Seasonally Dry Forest Species: Effects of Seed Traits and Population Environmental Conditions.

The persistence of subtropical seasonally dry forests urgently requires the implementation of ex situ conservation and restoration programs. We studied variation in seed traits and dormancy of six native species growing in seasonally dry Chaco forests of Argentina. We documented high intra- and interspecific variability in seed traits and dormancy. Fresh seeds of Geoffroea decorticans and Parasenegalia visco (Fabaceae) were water-permeable and nondormant (ND), while those of Parkinsonia praecox and Vachellia aroma (Fabaceae) were water-impermeable and had physical dormancy (PY). Seeds of Schnopsis lorentzii (Anacardiaceae) and Sarcomphalus mistol (Rhamnaceae) were water-permeable and had physiological dormancy (PD). Mechanical and chemical scarification were the most effective methods to break PY, and dry storage for 3 months was effective in breaking PD. Seeds of large-seeded species were ND or had PD, and those of small-seeded species had PY. Species inhabiting moist habitats had ND seeds, whereas those from seasonally dry habitats had seeds with PY or PD. These results suggest that seed traits and dormancy are species-specific and that intraspecific variation in seed traits is likely associated with high phenotypic plasticity of species in response to local environmental heterogeneity. These findings should be considered at the time of implementation of conservation techniques and for seed sourcing decisions for restoration.

Trade-offs between diaspore dispersal and dormancy within a spike of the invasive annual grass Aegilops tauschii.

MAIN CONCLUSION: Differences in dispersal and dormancy of heteromorphic diaspores of Aegilos tauschii may increase its flexibility to invade/occupy weedy unpredictable habitats by spreading risk in space and time. In plant species that produce dimorphic seeds, there often is a negative relationship between dispersal and dormancy, with high dispersal-low dormancy in one morph and low dispersal-high dormancy in the other, which may function as a bet-hedging strategy that spreads the risk of survival and ensures reproductive success. However, the relationship between dispersal and dormancy and its ecological consequences in invasive annual grasses that produce heteromorphic diaspores is not well studied. We compared dispersal and dormancy responses of diaspores from the basal (proximal) to the distal position on compound spikes of Aegilops tauschii, an invasive grass with heteromorphic diaspores. Dispersal ability increased and degree of dormancy decreased as diaspore position on a spike increased from basal to distal. There was a significant positive correlation between length of awns and dispersal ability, and awn removal significantly promoted seed germination. Germination was positively correlated with GA concentration and negatively correlated with ABA concentration, and the ABA: GA ratio was high in seeds with low germination/high dormancy. Thus, there was a continuous inverse-linear relationship between diaspore dispersal ability and degree of dormancy. This negative relationship between diaspore dispersal and degree of dormancy at different positions on a spike of Aegilops tauschii may facilitate seedling survival in space and time.

Burning lignin: overlooked cues for post-fire seed germination.

Information about smoke cues for seed germination is fundamental to understanding fire adaptation. Recently, lignin-derived syringaldehyde (SAL) was identified as a new smoke cue for seed germination, which challenges the assumption that cellulose-derived karrikins are the primary smoke cues. We highlight the overlooked association between lignin and the fire adaptation of plants.

Light controls alpine meadow community assembly during succession by affecting species recruitment from the seed bank.

Some research indicates that soil seed banks can promote species coexistence through storage effects. However, the seed bank mechanism that maintains plant assembly and its role in degraded grassland restoration are still not clear. We collected seed bank samples from early, mid and late secondary successional stages of an abandoned subalpine meadow on the Tibetan Plateau, and samples from each stage were exposed to full (i.e., natural), mid, and low light treatments in the field to represent light availability at the bottom/understory (soil surface) of a plant community in the early, mid and late stages of succession, respectively. Species richness, seed density, species composition, and community weighted mean values (CWMs) of seed mass of the species whose seeds germinated in soil samples were evaluated. In response to the light treatments, species richness increased significantly with increased light only for the late successional stage, seed density increased significantly with increased light only in the early and mid successional stages, and seed mass decreased significantly with increased light only in the mid and late successional stages. Species composition differed significantly among the light treatments only in the late successional stage. For the successional series, species richness and seed mass of the species that germinated increased significantly with succession only under mid and full light treatments. Seed density decreased significantly with succession in each light treatment. Species composition differed significantly between the early- and late stage and between the mid and late stage in each light treatment. Both the abiotic (light) and biotic (seed mass) factors influence seed bank recruitment to the plant community. Regeneration of small-seeded species in the seed bank was inhibited under low light in the late successional stage. The balance of stochastic and deterministic processes along a successional gradient was determined by regeneration from the seed bank depending on light intensity change. Differences in seed response to light intensity change largely determined plant community assembly. Our findings should help in the development of effective conservation and restoration strategies.

Exogenous gibberellin can effectively and rapidly break intermediate physiological dormancy of Amsonia elliptica seeds.

Accelerated global warming is leading to the loss of plant species diversity, and ex situ preservation of seeds is becoming an increasingly important aspect of species conservation. However, information on dormancy and germination is lacking in many endangered species. Amsonia elliptica (Apocynaceae) is the only Amsonia species native to Korea, and the South Korean Ministry of Environment has designated it Class II endangered wildlife. Nevertheless, the dormancy class and the dormancy breaking method for seeds of this species for germination are not precisely known. We identified the structure of A. elliptica seeds and the causes of dormancy, which inhibits germination. In addition, we tried to develop an effective germination promotion method by testing the wet stratified condition, which breaks dormancy, and the form of gibberellin that can replace it. Fresh seeds of A. elliptica imbibe water, but the covering layers (endosperm and seed coat) inhibit germination by mechanically restricting the embryo. Initial germination tests confirmed low embryo growth potential and physiological dormancy (PD). Restriction due to the covering layer was eliminated by seed scarification, and abnormal germination was observed. After 12 weeks of cold moist stratification at 4°C, only 12% of seeds germinated. However, 68.8% of seeds subjected to 8 weeks of warm moist stratification followed by 12 weeks of cold stratification germinated, indicating that warm stratification pretreatment before cold stratification is effective in breaking dormancy. A. elliptica seeds exhibited intermediate PD. Furthermore, 61.3% of seeds soaked in 500 mg/L GA4+7 for 14 days and incubated at 25/15°C germinated. Therefore, GA4+7 rapidly broke the dormancy of A. elliptica seeds compared with warm plus cold stratification treatment, thus providing an efficient method for seedling production.

Seed dormancy and germination of the endangered exceptional Hawaiian lobelioid Brighamia rockii.

Premise: The Campanulaceae (Lobelioideae) is the Hawaiian plant family with the most endangered and extinct species. Although seeds of Hawaiian lobelioids are desiccation tolerant, the species are exceptional (i.e., they present challenges at various stages of the conventional ex situ conservation chain) due to their generally poor seed survival at the conventional seed-banking temperature (-18°C). Both morphological dormancy (MD) and morphophysiological dormancy (MPD) have been identified in the seeds of other Hawaiian lobelioids; however, the class of dormancy and germination requirements of the Critically Endangered genus Brighamia have not yet been determined. Methods: We measured the embryonic growth in 12-week-old seeds of B. rockii and tested their germination at three temperature regimes (15/5°C, 20/10°C, and 25/15°C) in light and at 25/15°C in darkness. Results: The embryos grew prior to radicle emergence, and the seeds germinated rapidly to high percentages in all tested conditions. Discussion: Whether fresh B. rockii seeds have MD or MPD still needs to be determined; nevertheless, 12-week-old seeds germinated well in light and darkness, and thus the seeds can be used for conservation purposes. Germination in the dark suggests that the species may not form a long-lived soil seed bank in its native habitat.

Seed germination thermal niche differs among nine populations of an annual plant: A modeling approach.

Germination timing is an important determinant of survival and niche breadth of plants. The annual plant Nigella sativa occurs in diverse environments along a steep temperature gradient and thus is a suitable model for the study of germination behavior in response to temperature. We used a modeling approach to compare the germination thermal niche of seeds of nine populations of N. sativa produced in a common garden. Germination time courses were obtained by a newly developed process-based model, and thermal niche was visualized by plotting germination breadth as a function of after-ripening time. Seeds were sampled five times: immature (2 weeks before maturity), mature, and afterripened for 1, 2, and 5 months. Immature and mature seeds had a greater depth of dormancy than afterripened seeds, as estimated by lower values of high-limit temperatures (T h). Afterripening increased germination percentage, synchrony, and thermal niche breadth of all nine populations. The highest asynchrony was for immature and mature seeds, and afterripening enhanced synchrony. Based on the new graphical method, N. sativa has Type 1 nondeep physiological dormancy, and thus, the germination niche is narrow at seed maturity, leading to a delayed germination strategy that is highly dependent on thermal time accumulated during afterripening. Our findings show that there is considerable variation in the germination thermal niche among populations. Temperature regimes in the natural habitats of N. sativa have played a significant role in shaping variation in thermal niche breadth for seed germination of this annual species. The models used in our study precisely predict germination behavior and thermal niche under different environmental conditions. The germination synchrony model also can estimate germination pattern and degree of dormancy during the year, suggesting a useful method for quantification of germination strategies.

Seed position in spikelet as a contributing factor to the success of the winter annual invasive grass Aegilops tauschii.

Seed position - dependent effects on seed dormancy/germination are well documented at the inflorescence/infructescence level, but less is known about seeds at different positions within a dispersal unit. For the invasive winter annual grass Aegilops tauschii, we quantified morphology, mass and dormancy/germination of seeds from basal (1), middle (2), and distal (3) positions in two spikelet types (Left and Right). We also investigated seedling emergence, survival, plant size and seed production of plants from seeds in different spikelet positions of two spikelet types under different soil nutrient and water conditions. We found that these seed, seedling and plant traits performed as mirror images between the Left and Right spikelet types. The middle seed was significantly the longest and had the maximum mass, while the basal seed was the shortest and had medium mass. Middle seeds had the highest increase in mass during imbibition and the highest germination percentages and rates, while basal seeds had the lowest. Seedling emergence and survival, plant size and seed production for each position of seeds were highest in the added fertilizer combined with regular watering treatment and lowest in the no fertilizer combined with natural moisture, while height of plants derived from the middle and the distal seeds was significantly higher than that of plants derived from the basal seeds under all soil nutrient and water conditions. Seedling survival, number of tillers per plant and seed production per plant from the middle and distal seeds were significantly lower than those from basal seeds under all soil nutrient and water treatments. The considerable variation in seedling emergence and survival, plant size and seed production between seeds in different positions in the spikelet results in much flexibility in all stages of the life cycle, thereby likely contributing to the invasiveness of A. tauschii.

Negative impacts of human disturbances on the seed bank of subalpine forests are offset by climatic factors.

Precipitation and temperature in the subalpine region have increased dramatically in recent decades due to global warming, and human disturbances have continued to impact the vegetation in the region. Seed bank plays an important role in population recovery, but there are few studies on the synergistic effects of human disturbances and climate change on seed bank. We analyzed the synergistic effects of human disturbances and climate change on seed bank samples from 20 sites in the subalpine coniferous forest region using grazing and logging as the disturbance intensity gradient and precipitation and temperature as climate variables. The species diversity of aboveground vegetation all changed significantly (p < 0.05) with precipitation, temperature and disturbance level, while the seed bank richness and density did not. Furthermore, the species composition of the seed bank varied significantly less than that of the aboveground vegetation at different levels of disturbance (p < 0.001). Thus, seed bank showed a strong buffering capacity against the risk of local extinction caused by environmental changes that shift the species composition and diversity of aboveground vegetation. In addition, soil and litter are important influences controlling seed bank density in subalpine forests, and the results of structural equation modelling suggest that both disturbance and climate change can indirectly regulate the seed bank by changing the physicochemical properties of soil and litter. We conclude that increases in precipitation and temperature driven by climate change can buffer the negative effects of disturbances on the seed bank.

Effect of Light Intensity on Morphology, Photosynthesis and Carbon Metabolism of Alfalfa (Medicago sativa) Seedlings.

To understand how light intensity influences plant morphology and photosynthesis in the forage crop alfalfa (Medicago sativa L. cv. Zhongmu 1), we investigated changes in leaf angle orientation, chlorophyll fluorescence, parameters of photosynthesis and expression of genes related to enzymes involved in photosynthesis, the Calvin cycle and carbon metabolism in alfalfa seedlings exposed to five light intensities (100, 200, 300, 400 and 500 µmol m-2 s-1) under hydroponic conditions. Seedlings grown under low light intensities had significantly increased plant height, leaf hyponasty, specific leaf area, photosynthetic pigments, leaf nitrogen content and maximal PSII quantum yield, but the increased light-capturing capacity generated a carbon resource cost (e.g., decreased carbohydrates and biomass accumulation). Increased light intensity significantly improved leaf orientation toward the sun and upregulated the genes for Calvin cycle enzymes, thereby increasing photosynthetic capacity. Furthermore, high light (400 and 500 µmol m-2 s-1) significantly enhanced carbohydrate accumulation, accompanied by gene upregulation and increased activity of sucrose and starch-synthesis-related enzymes and those involved in carbon metabolism. Together, these results advance our understanding of morphological and physiological regulation in shade avoidance in alfalfa, which would guide the identification of suitable spatial planting patterns in the agricultural system.

Plant Growth Regulators Improve Grain Production and Water Use Efficiency of Foeniculum vulgare Mill. under Water Stress.

The development of methods increasing plant water use efficiency (WUE) would enhance the ability to grow wild aromatic and medicinally important species. The aim of this research was to determine the effect of plant growth regulators (PGRs) applied by spraying on stress resistance and WUE of fennel subjected to water stress. Plants in the generative stage were more drought tolerant than those in the vegetative stage. Water stress at vegetative stage decreased plant biomass and grain yield by 60% and 61%, respectively. Severe water stress in vegetative stage reduced grain production by 56%, and grains had 43% lower mass than those from non-stressed plants. Application of PGRs at both stages of growth increased grain yield and biomass, but the magnitude of increase depended on the type and application time of PGRs. Plants grown in well-watered conditions and sprayed with methyl jasmonate during the vegetative stage had the highest grain production (2.7 g plant-1), whereas under moderate water stress, plants yielded the best (2.1 g plant-1) when sprayed with epibrassinolide. The maximum WUE for grain (0.91 g L-1) and essential oil production (20 mg L-1) was noted in plants exposed to moderate stress and treated with methyl jasmonate during the vegetative stage.

Germination of Seeds from Flowers along a Continuum of Long to Short Styles in the Cold Desert Perennial Herb Ixiolirion songaricum.

We compared seed set, mass, and dormancy/germination of seeds from flowers at three points on the style-length continuum [long (LS), intermediate (IS), and short (SS) styles] in Ixiolirion songaricum. The effects of open and hand pollination (self and cross with pollen from upper and lower-level stamens) on seed set, mass, and dormancy/germination were assessed. Most freshly-matured seeds from LS, IS, and SS flowers were dormant, and dormancy was broken under laboratory and field conditions. After-ripened seeds from LS and IS flowers germinated to significantly higher percentages than those from SS flowers. In all pollination treatments, seed set and mass were significantly higher for LS and IS than for SS flowers. Seed set, mass, and germination for LS, IS, and SS flowers were significantly higher in open-pollinated and in cross-pollinated with pollen from upper and lower-level stamens than in self-pollination with pollen from upper- and lower-level stamens. These differences in offspring reproductive traits may be adaptive for I. songaricum in its rainfall-unpredictable environment. This is the first study to demonstrate the association between style length and germination in a species with continuous variation in style length.