Pollen availability,seed production and seed predator clutch size in a tephritid—thistle system

Summary We develop a simple model explaining clutch size behaviour ofOrellia ruficauda on its principle host in North America,Cirsium arvense. Offspring of flies feed solely on thistle seeds and seed production is pollen-limited. Thus, female flies risk reduced offspring fitness when committing large clutches to hosts (female flower heads) occurring in localities where male plants are locally absent. We therefore predict that attacked hosts will contain fewer eggs in such localities, a prediction that is consistent with data obtained in the field: large clutches are never laid in flower heads in low-pollination localities. However, larvae reared from such low-quality hosts are significantly smaller on average and will therefore carry smaller egg loads as adults. Small clutches in poor-quality hosts may thus be an expression of lower per-adult fecundity. Nevertheless, sufficient numbers of large, fecund flies are produced in low-pollination localities to make this last explanation less convincing.     

Do seed and microsite limitation interact with seed size in determining invasion patterns in flooding Pampa grasslands?

Additive influences of the invasion ability of species (invasiveness) and the characteristics of the habitat which make it invasible (invasibility) cannot fully explain grassland invasion patterns. We tested the hypothesis that different species assemblages of grassland communities may partly result from interactive influences between the relative invasiveness of available species and community invasibility. During 10 months, we evaluated seed and microsite limitation of seedling emergence, survival, and recruitment of plants belonging to species with different seed size (large-seeded species vs. small-seeded species), with in a two-phase community mosaic typical of semi-natural grasslands in the southern flooding Pampa of Argentina. Seeds of large versus small-seeded species were sown either in species-poor patches dominated by a tall tussock grass (“pajonal”) or in species-rich patches dominated by short grasses (“matrix”), subjected to different levels of canopy disturbance (cut vs. uncut). Seed addition promoted seedling emergence for 7 out of the 10 species sown, and this effect was higher for large than for small-seeded species. After seed limitations were removed, interactive effects among seed size, community state and canopy disturbance reflected a strong positive influence of seed size on plant recruitment only in cut pajonal patches. Therefore, according to the stage of invasion process, relative species success may depend on non-interactive (seed-size effect on seed limitation to seedling emergence) or interactive influences among species invasibility and community invasiveness (from seedling emergence to plant recruitment). As a general conclusion, different assemblages of species are expected to successfully colonize spatially close grassland patches, according to both the available invasible species (seed size) and the community state and stage (species composition and canopy disturbance).     

Is fire a selective force of seed size in pine species?

Fire is selectively shaping most of the traits of plants growing in fire-prone environments. However, seed size and other features related to seed production have not been studied in the light of the evolutionary role of fire. Our research tests the hypothesis that larger seeds have a higher chance of surviving wildfires and produce more vigorous seedlings with a lower death rate. To test this hypothesis the germination and early seedling growth of five Spanish pine species were studied. Weight, length and width of all seeds were measured. The biomass (fresh and dry weight) and length (root and total) of subsequent seedlings were also measured after 30 d from emergence. Seeds were submitted to elevated temperatures for periods in which the chance of survival was 50 % (calculated by means of a logistic model for each pine species). The differences observed among species suggests that fire may be adaptively shaping seed size in pines with larger seeds (Pinus canariensis and P. pinaster), because larger seeds are more likely to survive after heat shocks. Furthermore, in P. canariensis, seedlings after heat treatment are even larger than those submitted to control. In P. halepensis, despite being well adapted to fire, our results indicated no relationships between fire and seed characteristics. Finally, although heat treatment has a general adverse effect on seedling growth in the case of the two subalpine pines, we have detected a positive relationship between seed size and seedling growth but only in the largest seeds. This might also suggest the relevance of fire as a selective force for these pines which is outperformed by the relevance of dispersal and emergence time as adaptive traits in the post-fire scenario.     

Why are small seeds dispersed through animal guts: large numbers or seed size per se?

Long-distance dispersal of seeds is an important process in metapopulation dynamics and in plant migrations, but at the same time extremely difficult to observe or quantify directly. If seed dispersal ability were related to attributes of seeds or motherplants, long-distance seed dispersal would be predictable by indirect approximation using easy-to-measure traits. Seed size has been suggested to be such a key trait in seed dispersal ability. However, having smaller seeds also implies having more numerous seeds per plant individual (given equal reproductive effort), and consequently increases the probability of seeds being ingested accidentally. The question is whether small-seeded species are more abundant in herbivore dung because smaller seed size increases survival rate during gut passage or because they are produced (and ingested) in greater numbers than larger seeds. We investigated endozoochorous seed dispersal via cattle grazing a meadow, and related seed abundance in dung samples to seed attributes. We found that seeds were ingested and passed through the bovine intestinal tract in proportion to the numbers produced per unit area in the grazed vegetation. In contrast, no relationship could be found between endozoochorous dispersal potential (measured as abundance of seeds in dung samples corrected for seed output in the grazed vegetation) and seed attributes such as seed mass, seed shape (roundness), and thickness of the seed coat. This finding underlines the importance of seed number in plant dispersal ability. In addition, it shows that grazing mammals may constitute an important dispersal vector for many plant species conventionally classified as 'unspecialised'.     

Geographic variation for elaiosome–seed size ratio and its allometric relationship in two closely related Corydalis species

Abstract

Background: Elaiosomes serve as a reward for seed-dispersing ants. The size of the elaiosome and the elaiosome–seed mass (ESM) ratio are expected to affect seed dispersal, whilst seed size affects the establishment of seedlings. Elaiosomes and seeds can undergo independent change, thus variation in these traits may arise through heterogeneous selection for seedling establishment and for dispersal. Only a few studies have examined intraspecific variation in these traits.

Aims: The aim of this study was to determine if the ratio of elaiosome–seed mass was different in two co-occurring congeneric plant species (Corydalis intermedia and C. solida). Under the hypothesis that the ESM ratio affected diaspore attractiveness to ants, the allometric relationship between elaiosome and seed mass was used to infer how selection by ants may shape the diaspore.

Methods: Seed and elaiosome mass were measured in fruits collected from central and marginal populations of the two plant species. The allometric relationship between seed and elaiosome mass was obtained using major axis regression. In situ cafeteria experiments estimated the removal rate of diaspores by ants.

Results: Intra- and interspecific variation in diaspore traits were found. The selfing C. intermedia produced heavier seeds and elaiosomes than the outcrossing C. solida. Ants removed more diaspores from C. intermedia where both species were present. A slope larger than one characterised the allometric relationship between elaiosome and seed mass in both species. This slope was twice as steep in the central populations of C. intermedia compared to marginal ones.

Conclusions: Results indicate that in C. intermedia elaiosome mass must increase more than proportionally with increasing seed mass for the diaspore to remain attractive to ants. The direction of interspecific differences suggests that a plant-mating system may affect selection for dispersal.     

Bridging meta-analysis and the comparative method: a test of seed size effect on germination after frugivores’ gut passage

Most studies using meta-analysis try to establish relationships between traits across taxa from interspecific databases and, thus, the phylogenetic relatedness among these taxa should be taken into account to avoid pseudoreplication derived from common ancestry. This paper illustrates, with a representative example of the relationship between seed size and the effect of frugivores gut on seed germination, that meta-analytic procedures can also be phylogenetically corrected by means of the comparative method. The conclusions obtained in the meta-analytical and phylogenetical approaches are very different. The meta-analysis revealed that the positive effects that gut passage had on seed germination increased with seed size in the case of gut passage through birds whereas decreased in the case of gut passage through non-flying mammals. However, once the phylogenetic relatedness among plant species was taken into account, the effects of gut passage on seed germination did not depend on seed size and were similar between birds and non-flying mammals. Some methodological considerations are given to improve the bridge between the meta-analysis and the comparative method.     

Why is there variation in mean seed size among plants within single populations? test of the fertilization efficiency hypothesis

We tested the fertilization efficiency hypothesis, which attempts to explain mean seed size variation among plants within single populations, by comparing the patterns of seed size variation between chasmogamous (CH) flowers and cleistogamous (CL) flowers in Impatiens noli-tangere and Viola grypoceras, respectively. The fertilization efficiency hypothesis predicts that larger plants produce larger seeds if the number of pollen grains captured by a plant increases with increased allocation of resources to its attractive structures (e.g., corolla and nectar), but with diminishing gains. Thus, seed size should depend on plant size in seeds from CH flowers because of the diminishing gains of capturing pollen in these flowers, whereas seed size should not depend on plant size in seeds from CL flowers because CL flowers need not capture outcross pollen. We found significant positive correlations between mean seed size per plant and plant size for seeds from CH flowers in both species. However, there was no significant positive correlation between these two factors for seeds from CL flowers of both species. The results of the present investigations were thus consistent with the fertilization efficiency hypothesis.     

Effects of topographic position,leaf litter and seed size on seedling demography in a semi-deciduous tropical forest in Panamá

This study examined whether topography-induced gradients in water potential and leaf litter depth contribute to species coexistence in tropical forests through species-specific effects on seedling emergence and mortality. Seedling emergence and mortality were followed for a period of 12 months in 36 (1 × 2 m) plots on Barro Colorado Island (BCI), Panamá. Plots with and without litter were distributed on slope and plateau sites in three catchments. In the absence of manipulations, the lower litter depth on slopes resulted in approximately four times as many emergent seedlings than on plateau sites. However, litter depth had little effect on seedling community composition. By the end of the first dry-season, post-emergence, there were no significant differences in surviving seedling numbers between any treatments. There were differences in the emergent seedling community between slope and plateau sites within the same catchment as well as differences in composition between catchments, suggesting that both niche partitioning and dispersal limitation might play a role in structuring seedling community composition. During the wet-season seedling mortality was highest on slope sites although this pattern was reversed during the dry-season. In both seasons mortality was higher for small-seeded species. These results demonstrate that gradients in water potential related to topography impact on patterns of seedling emergence and mortality although processes in the first year after emergence may be insufficient to explain observed habitat preferences of adult plants.     

Are plant populations seed limited? A critique and meta-analysis of seed addition experiments

We examine the relative importance of processes that underlie plant population abundance and distribution. Two opposing views dominate the field. One posits that the ability to establish at a site is determined by the availability of suitable microsites (establishment limitation), while the second asserts that recruitment is limited by the availability of seeds (seed limitation). An underlying problem is that establishment and seed limitation are typically viewed as mutually exclusive. We conducted a meta-analysis of seed addition experiments to assess the relative strength of establishment and seed limitation to seedling recruitment. We asked (1) To what degree are populations seed and establishment limited? (2) Under what conditions (e.g., habitats and life-history traits) are species more or less limited by each? (3) How can seed addition studies be better designed to enhance our understanding of plant recruitment? We found that, in keeping with previous studies, most species are seed limited. However, the effects of seed addition are typically small, and most added seeds fail to recruit to the seedling stage. As a result, establishment limitation is stronger than seed limitation. Seed limitation was greater for large-seeded species, species in disturbed microsites, and species with relatively short-lived seed banks. Most seed addition experiments cannot assess the relationship between number of seeds added and number of subsequent recruits. This shortcoming can be overcome by increasing the number and range of seed addition treatments.     

Non‐native conditions favor non‐native populations of invasive plant: demographic consequences of seed size variation?

Trait differences between native and non‐native populations may explain the greater abundance and impact of some organisms in their non‐native ranges than in their native ranges. Here, we conducted reciprocal common gardens in southwestern Turkey (home) and central Argentina (away) to explore the hypothesis that the greater success of the invasive ruderal Centaurea solstitialis in Argentina than Turkey is partially explained by differences between home and away populations. Unusual among common gardens, our experimental design included seed additions to explicitly evaluate population level responses, as well as disturbance and no‐disturbance treatments. We documented seed mass in native and non‐native populations, and during the experiment, we periodically measured density, plant size, and herbivory. After six months, we determined the establishment of plants for populations from both origins in both home and away common gardens. Seed mass was two times larger for Argentinean than Turkish populations. Density, plant size and final establishment were also greater for plants from Argentinean than from Turkish populations, but only in the common garden in Argentina. In Turkey, no differences between population origins were detected for these variables. Herbivory was similar for populations from both origins in both common gardens. As expected, disturbance generally increased plant performance in both regions. Our results suggest that increased seed size in non‐native populations may have demographic consequences under non‐native conditions that can contribute to the invasive success of C. solstitialis. This is the first reciprocal common garden that supports the idea that seed size variation contributes to demographic differences for an invasive species between native and non‐native distributions, but our findings further suggest that seed size effects on demography depend on the ecological context in which population processes occur.     

Does size matter?

For 40 years, the debate has raged. Do mammalian cells monitor cell size when deciding whether to divide? More recent models suggest an indirect solution, but the field is far from reaching a final verdict.     

Do large‐seeded herbs have a small range size? The seed mass–distribution range trade‐off hypothesis

We aimed to introduce and test the “seed mass–distribution range trade‐off” hypothesis, that is, that range size is negatively related to seed mass due to the generally better dispersal ability of smaller seeds. Studying the effects of environmental factors on the seed mass and range size of species, we also aimed to identify habitats where species may be at risk and need extra conservation effort to avoid local extinctions. We collected data for seed mass, global range size, and indicators for environmental factors of the habitat for 1,600 species of the Pannonian Ecoregion (Central Europe) from the literature. We tested the relationship between species’ seed mass, range size, and indicator values for soil moisture, light intensity, and nutrient supply. We found that seed mass is negatively correlated with range size; thus, a seed mass–distribution range trade‐off was validated based on the studied large species pool. We found increasing seed mass with decreasing light intensity and increasing nutrient availability, but decreasing seed mass with increasing soil moisture. Range size increased with increasing soil moisture and nutrient supply, but decreased with increasing light intensity. Our results supported the hypothesis that there is a trade‐off between seed mass and distribution range. We found that species of habitats characterized by low soil moisture and nutrient values but high light intensity values have small range size. This emphasizes that species of dry, infertile habitats, such as dry grasslands, could be more vulnerable to habitat fragmentation or degradation than species of wet and fertile habitats. The remarkably high number of species and the use of global distribution range in our study support our understanding of global biogeographic processes and patterns that are essential in defining conservation priorities.     

Does the seed size/number trade‐off model determine plant community structure? An assessment of the model mechanisms and their generality

This paper examines four key mechanisms of the seed size/number trade-off (SSNT) models to assess their relevance to a general understanding of plant community structure. Mechanism 1 is that large seeds have a greater probability of winning in competition against smaller seeds. I provide interspecific experimental evidence that there is a competitive hierarchy among seedlings based on seed size. Mechanism 2 is that a trade-off exists between the number and size of seeds produced for a given reproductive allocation. Negative correlations between seed size and number were found consistently across a range of species from a range of habitats, from published literature. Mechanism 3, that seedling-seedling competition is an important influence on species composition, was found to exist potentially in a range of environments, including annual-dominated, post-fire and gap-dynamic communities. However, there is little quantitative evidence available and this is likely to be a restrictive mechanism. Mechanism 4, that small seeds are superior colonists due to their greater number, was tested in a field experiment in a calcareous grassland community. No supporting evidence was found, suggesting that the SSNT is not an important determinant of structure in this community. Thus two of the four mechanisms can be considered to hold true generally, while the third mechanism may be valid in particular environments. The fourth mechanism did not apply in the community tested, but could be tested in a wider range of communities.     

Evaluation of seed storage-protein gene 5′ untranslated regions in enhancing gene expression in transgenic rice seed

5′ untranslated regions (UTRs) are important sequence elements that modulate the expression of genes. Using the β-glucuronidase (GUS) reporter gene driven by the GluC promoter for the rice-seed storage-protein glutelin, we evaluated the potential of the 5′-UTRs of six seed storage-protein genes in enhancing the expression levels of the foreign gene in stable transgenic rice lines. All of the 5′-UTRs significantly enhanced the expression level of the GluC promoter without altering its expression pattern. The 5′-UTRs of Glb-1 and GluA-1 increased the expression of GUS by about 3.36- and 3.11-fold, respectively. The two 5′-UTRs downstream of the Glb-1, OsAct2 and CMV35S promoters also increased GUS expression level in stable transgenic rice lines or in transient expression protoplasts. Therefore, the enhancements were independent of the promoter sequence. Real-time quantitative RT-PCR analysis showed that the increase in protein production was not accompanied by alteration in mRNA levels, which suggests that the enhancements were due to increasing the translational efficiencies of the mRNA. The 5′-UTRs of Glb-1 and GluA-1, when combined with strong promoters, might be ideal candidates for high production of recombinant proteins in rice seeds.     

Roles of the Arabidopsis G protein γ subunit AGG3 and its rice homologs GS3 and DEP1 in seed and organ size control

The size of seeds and organs is coordinately determined by cell proliferation and cell expansion, but the mechanisms that set final seed and organ size are largely unknown in plants. In a recent study, we have demonstrated that the plant specific G protein γ subunit (AGG3) promotes seed and organ growth by increasing the period of proliferative growth in Arabidopsis. AGG3 is localized in plasma membrane and interacts with the G protein β subunit (AGB1). Homologs of AGG3 in rice (GS3 and DEP1/qPE9–1) have been identified as important quantitative trait loci for seed size and yield. However, rice GS3 and DEP1 influence seed and organ growth by restricting cell proliferation. Here, we discuss the possible molecular mechanisms by which Arabidopsis AGG3 and its rice homologs GS3 and DEP1 control seed and organ size.