Trade-offs between seed dispersal and dormancy in an amphi-basicarpic cold desert annual.

BACKGROUND AND AIMS: Several studies have demonstrated trade-offs between depth of seed dormancy and dispersal ability for diaspore-dimorphic species. However, relatively little is known about trade-offs between these two life history traits for a species that produces more than two diaspore morphs. The aim of this study was to investigate the relationship between seed dormancy and dispersal in Ceratocarpus arenarius, an amphi-basicarpic cold desert annual that produces a continuum of dispersal unit morphs. METHODS: A comparison was made of dispersal and dormancy breaking/germination responses of dispersal units from ground level (a), the middle of the plant canopy (c) and the top of the plant canopy (f). Various features of the morphology and mass of dispersal units and fruits (utricles) were measured. The role of bracteoles in diaspore dispersal by wind, settlement onto the soil surface and dormancy/germination was determined by comparing responses of intact dispersal units and fruits. Movement of dispersal units by wind and animals, seed after-ripening, germination phenology and the presence of water-soluble germination inhibitors in bracteoles were tested using standard procedures. KEY RESULTS: Dispersal units a, c and f differed in morphology and mass; in the majority of cases, extremes were exhibited by a and f, with c being intermediate. Overall, relative dispersal ability was f > c > a, whereas relative intensity of dormancy was a > c > f. Bracteoles increased dispersal distance by wind, enhanced settlement of diaspores onto the soil surface and mechanically inhibited germination. CONCLUSIONS: The results provide evidence for a model in which there is a continuous inverse-linear relationship between diaspore dispersal ability and depth of dormancy. Thus, dispersal unit heteromorphism of C. arenarius results in a continuum, from no dispersal ability/high dormancy (dispersal unit a) to high dispersal ability/low dormancy (unit f), which may be a bet-hedging strategy in the cold desert environment.

Two-dimensional chromatographic analysis using three second-dimension columns for continuous comprehensive analysis of intact proteins.

We develop a new two-dimensional (2D) high performance liquid chromatography (HPLC) approach for intact protein analysis. Development of 2D HPLC has a bottleneck problem - limited second-dimension (second-D) separation speed. We solve this problem by incorporating multiple second-D columns to allow several second-D separations to be proceeded in parallel. To demonstrate the feasibility of using this approach for comprehensive protein analysis, we select ion-exchange chromatography as the first-dimension and reverse-phase chromatography as the second-D. We incorporate three second-D columns in an innovative way so that three reverse-phase separations can be performed simultaneously. We test this system for separating both standard proteins and E. coli lysates and achieve baseline resolutions for eleven standard proteins and obtain more than 500 peaks for E. coli lysates. This is an indication that the sample complexities are greatly reduced. We see less than 10 bands when each fraction of the second-D effluents are analyzed by sodium dodecyl sulfate - polyacrylamide gel electrophoresis (SDS-PAGE), compared to hundreds of SDS-PAGE bands as the original sample is analyzed. This approach could potentially be an excellent and general tool for protein analysis.

Effect of Seed Position on Parental Plant on Proportion of Seeds Produced with Nondeep and Intermediate Physiological Dormancy.

The position in which seeds develop on the parental plant can have an effect on dormancy-break and germination. We tested the hypothesis that the proportion of seeds with intermediate physiological dormancy (PD) produced in the proximal position on a raceme of Isatis violascens plants is higher than that produced in the distal position, and further that this difference is related to temperature during seed development. Plants were watered at 3-day intervals, and silicles and seeds from the proximal (early) and distal (late) positions of racemes on the same plants were collected separately and tested for germination. After 0 and 6 months dry storage at room temperature (afterripening), silicles and seeds were cold stratified for 0-16 weeks and tested for germination. Mean daily maximum and minimum temperatures during development/maturation of the two groups of seeds did not differ. A higher proportion of seeds with the intermediate level than with the nondeep level of PD was produced by silicles in the proximal position than by those in the distal position, while the proportion of seeds with nondeep PD was higher in the distal than in the proximal position of the raceme. The differences were not due only to seed mass. Since temperature and soil moisture conditions were the same during development of the seeds in the raceme, differences in proportion of seeds with intermediate and nondeep PD are attributed to position on parental plant. The ecological consequence of this phenomenon is that it ensures diversity in dormancy-breaking and germination characteristics within a seed cohort, a probable bet-hedging strategy. This is the first demonstration of position effects on level of PD in the offspring.

Effects of germination season on life history traits and on transgenerational plasticity in seed dormancy in a cold desert annual.

The maternal environment can influence the intensity of seed dormancy and thus seasonal germination timing and post-germination life history traits. We tested the hypotheses that germination season influences phenotypic expression of post-germination life history traits in the cold desert annual Isatis violascens and that plants from autumn- and spring-germinating seeds produce different proportions of seeds with nondeep and intermediate physiological dormancy (PD). Seeds were sown in summer and flexibility in various life history traits determined for plants that germinated in autumn and in spring. A higher percentage of spring- than of autumn-germinating plants survived the seedling stage, and all surviving plants reproduced. Number of silicles increased with plant size (autumn- > spring-germinating plants), whereas percent dry mass allocated to reproduction was higher in spring- than in autumn-germinating plants. Autumn-germinating plants produced proportionally more seeds with intermediate PD than spring-germinating plants, while spring-germinating plants produced proportionally more seeds with nondeep PD than autumn-germinating plants. Flexibility throughout the life history and transgenerational plasticity in seed dormancy are adaptations of I. violascens to its desert habitat. Our study is the first to demonstrate that autumn- and spring-germinating plants in a species population differ in proportion of seeds produced with different levels of PD.

Seed Germination Ecology of the Cold Desert Annual Isatis violascens (Brassicaceae): Two Levels of Physiological Dormancy and Role of the Pericarp.

The occurrence of various species of Brassicaceae with indehiscent fruits in the cold deserts of NW China suggests that there are adaptive advantages of this trait. We hypothesized that the pericarp of the single-seeded silicles of Isatis violascens restricts embryo expansion and thus prevents germination for 1 or more years. Thus, our aim was to investigate the role of the pericarp in seed dormancy and germination of this species. The effects of afterripening, treatment with gibberellic acid (GA3) and cold stratification on seed dormancy-break were tested using intact silicles and isolated seeds, and germination phenology was monitored in an experimental garden. The pericarp has a role in mechanically inhibiting germination of fresh seeds and promotes germination of nondormant seeds, but it does not facilitate formation of a persistent seed bank. Seeds in silicles in watered soil began to germinate earlier in autumn and germinated to higher percentages than isolated seeds. Sixty-two percent of seeds in the buried silicles germinated by the end of the first spring, and only 3% remained nongerminated and viable. Twenty to twenty-five percent of the seeds have nondeep physiological dormancy (PD) and 75-80% intermediate PD. Seeds with nondeep PD afterripen in summer and germinate inside the silicles in autumn if the soil is moist. Afterripening during summer significantly decreased the amount of cold stratification required to break intermediate PD. The presence of both nondeep and intermediate PD in the seed cohort may be a bet-hedging strategy.

Germination season and watering regime, but not seed morph, affect life history traits in a cold desert diaspore-heteromorphic annual.

Seed morph, abiotic conditions and time of germination can affect plant fitness, but few studies have tested their combined effects on plasticity of plant life history traits. Thus, we tested the hypothesis that seed morph, germination season and watering regime influence phenotypic expression of post-germination life history traits in the diaspore-heteromorphic cold desert winter annual/spring ephemeral Diptychocarpus strictus. The two seed morphs were sown in watered and non-watered plots in late summer, and plants derived from them were watered or not-watered throughout the study. Seed morph did not affect phenology, growth and morphology, survival, dry mass accumulation and allocation or silique and seed production. Seeds in watered plots germinated in autumn (AW) and spring (SW) but only in spring for non-watered plots (SNW). A high percentage of AW, SW and SNW plants survived and reproduced, but flowering date and flowering period of autumn- vs. spring-germinated plants differed. Dry mass also differed with germination season/watering regime (AW > SW > SNW). Number of siliques and seeds increased with plant size (AW > SW > SNW), whereas percent dry mass allocated to reproduction was higher in small plants: SNW > SW > AW. Thus, although seed morph did not affect the expression of life history traits, germination season and watering regime significantly affected phenology, plant size and accumulation and allocation of biomass to reproduction. Flexibility throughout the life cycle of D. strictus is an adaptation to the variation in timing and amount of rainfall in its cold desert habitat.