Seed yield,seed size and germination behaviour in the annualPogogyne abramsii

Summary Models of the evolution of seed dormancy reveal that dormancy is favoured either when opportunities for establishment vary over time and when there is wide variation in the probability of success, or when the probability of success is limited by frequency dependence. Empirical evidence supporting the temporal heterogeneity hypothesis exists, but there is scant evidence for dormancy being favoured by frequency dependent competition among seedlings. We test the hypothesis that the intensity of between-sib competition should favour a positive relationship between maternal fecundity and seed dormancy. This hypothesis is supported for the rare, vernal pool annual,Pogogyne abramsii: the proportion of dormant offspring was significantly higher among high fecundity mothers than among other mothers. Dormancy inP. abramsii is controlled by the seed coat, a maternal tissue, so delaying germination favours the inclusive fitness of mothers by reducing the potential for competition among siblings. Seed weight and time to first germination varied significantly amongP. abramsii plants and mean seed weight increased linearly with plant biomass. Seed weight and seed number are independently regulated by plant size. Overall, seed weight varied 10-fold and variability in seed weight within mothers was not explained by plant biomass, seed yield or mean seed weight. GerminableP. abramsii seeds were significantly heavier than dormant seeds, and germinable seeds heavier than 0.20 mg germinated more rapidly than those smaller than 0.20 mg.     

Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination

Canopy gap and chilling requirements for seed germination were assessed in 61 and 44 species, respectively, in the temperate rain forest of southern Chile. Germination assays within canopy gap and understorey were carried out under natural conditions. Germination tests in cold stratified and non-stratified seeds were performed under laboratory conditions. Seeds were collected of common trees, shrubs, vines and perennial herbs of forests in southern Chile. Final percent germination was significantly enhanced under canopy gap conditions in 19 species, and significantly reduced in 11 species. Germination proved indifferent under gap vs. understorey conditions in half the species tested. Cold stratification affected germination in 11 species: significantly increasing final germination of four species, and significantly decreasing final percent germination of seven species. A Principal Components Analysis (PCA) was applied with the purpose of identifying groups of species with similar germination strategies. Four attributes were included in the PCA: final germination percentage in canopy gap, germination rate in the laboratory, and gap-and chilling-dependency indices (EGAP and STRAT, respectively). The first axis separated species mainly on EGAP variation while the second axis separated them mainly according to STRAT variation; the two axes together explaining 73% of the among-species variation. A small group of trees and vines germinating best in the understorey and neutral to chilling could clearly be distinguished from the remaining species analyzed. Multifactorial ANOVAs were used for evaluating the combined effect of successional status, seed mass, dispersal period, life form, phylogenetic categories, and dispersal syndrome on EGAP and STRAT variation. The EGAP value of secondary successional species proved significantly greater than that of primary successional species and species with endozoochorous seed dispersal were significantly less dependent on chilling (according to STRAT values) compared to species with mainly wind-dispersed seeds. The possible implications of these results for seedling establishment are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Seed mass and dormancy of annual plant populations and communities decreases with aridity and rainfall predictability

Several theoretical and empirical studies have examined the influence of environmental conditions on seed traits and germination strategies of annual species. A positive relationship between seed mass and dormancy has been described for annuals occupying climatically unpredictable ecosystems. Larger-seeded species tend to have higher seedling survival rates, while dormancy allows a bet-hedging strategy in unpredictable environments. Until now, these ideas have been addressed primarily for only one or a few focal species, without considering differences among populations and communities. The novelty of the present study lies in the population and community-level approach, where a comprehensive seed trait database including 158 annual species occurring along a gradient of rainfall variability and aridity in Israel was used to ask the following question: Does average seed mass and dormancy of annual populations and communities decrease with increasing aridity and rainfall unpredictability?Soil seed bank samples were collected at the end of the summer drought, before the onset of the rains, from four plant communities. Germination was tested under irrigated conditions during three consecutive germination seasons to determine the overall seed germinability in each soil sample. Seed mass was obtained from newly produced seeds collected at the study sites in late spring. The community level results showed that, in contrast to common theoretical knowledge, seed mass and dormancy of the dominant annual species decreased with increasing aridity and rainfall variability. Accordingly, a negative correlation was found between seed mass and seed germination fractions. The present study demonstrates that an analysis of seed traits along climatic gradients is significantly improved by approaches that target both population and community levels simultaneously. A critical evaluation sheds new light upon the selective pressures that act on seed ecology of annuals along a climatic gradient and facilitates formulation of more mechanistic hypotheses about factors governing critical seed traits.     

Seed size and shade-tolerance: a comparative analysis of North American temperate trees

Large seed size is a trait associated with plant species of mature, closed habitats, and is thought to supply an ample nutrient reserve necessary for seedling establishment. While this relationship has been shown for annuals and short-lived perennials, it is poorly documented for trees. A comparative method was used to determine whether North American temperate tree species which typically establish in shady conditions have larger seeds than those requiring more open conditions. Both angiosperms (hardwoods) and gymnosperms (conifers) were involved in the study. A significant relationship was found between large seed mass and shade-tolerance for angiosperms, but not for gymnosperms. These contrasting results seem to relate to unique evolutionary opportunities or constraints in different taxa. The absence of a relationship for gymnosperms is discussed in terms of character traits which might prevent seed size selection in response to shade. The finding for angiosperms underlines the importance of the regeneration niche in promoting coexistence among temperate hardwood species through seed size differentiation. Received: 18 March 1997 / Accepted: 26 November 1997     

The ecology of very small-seeded shade-tolerant trees and shrubs in lowland rain forest in Singapore

A study was made in Bukit Timah Nature Reserve of niche differentiation among 11 woody species that have very small seeds (22–460 µg) and establish in deep shade, i.e., where the indirect site factor (isf) under cloudy conditions is 2%, and one taxonomically related light-demander (seed mass 33 µg). Comparative observations were made on changing light requirement with age. The species varied from shrubs and treelets (Urophyllum) via small to medium-height trees (Ficus, Pternandra) to tall trees (Gynotroches and Pellacalyx). In general, the shorter species fruited in deep shade while the taller needed direct irradiance. All produced ripe fruit at least once a year.In 1–3% isf seedling mortality over 11 months was 24% for Urophyllum hirsutum and 57% for Pternandra echinata (two strongly shade-tolerant species); survivors produced appreciable new stem and leaf tissue. For two more light-demanding species (Ficus grossularioides and F. fistulosa) mean lamina area was much larger in small gaps (10% isf) than in the understorey (<3% isf), while for Urophyllum hirsutum it was much smaller. These three species showed the greatest difference in SLA. There were few significant intraspecific differences between gaps and understorey in concentrations of N, P or K. In a valley where there was no significant difference in isf between a path and adjacent understorey, seedlings of species with seeds <1 mg mean dry mass were five times as dense on transects along steep pathside banks as in the understorey 5 m upslope. The mean mid-bank slope was 66° (cf. 20° upslope). On mid-bank transects litter had a cover of 36% (cf. 95% upslope), and was thinner (0.8 vs 3.2 cm). The soil at 0–7 cm depth into the mid-bank had much lower concentrations of organic matter, total N and P (but not total K) and fine roots than that of upslope transects. All species were found to have vesicular-arbuscular mycorrhiza when 1–2 m tall in deep shade, several having very dense infections. Seedling form and responses are discussed in relation to potential adaptive value and possible evolutionary origin.     

Effects of altitude and seed size on germination and seedling survival of heathland plants in north Spain

The effects of altitude and seed size on germination and seedling survival were studied in Calluna vulgaris, Erica cinerea and Erica vagans. Experiments were carried out in the laboratory over a one year period. Seeds collected from heathlands of different altitudes were divided in two size classes. They were sowed on moist filter paper inside Petri dishes which were placed in chambers at 20 °C and a photoperiod of 12 h light/12 h darkeness. The seeds of Calluna vulgaris were the first to begin germination and had the highest rate and percentage of germination. The germination of Erica vagans was moderate, while seeds of Erica cinerea germinated later and the germination was very low. Seeds of Calluna vulgaris and E. cinerea collected at the highest altitudes had the highest germination percentages. Seed size in Calluna vulgaris and E. cinerea did not affect germination. However, large seeds of Erica vagans had higher germination rates and percentages than small seeds. The large seed size of Calluna vulgaris contributes to a better survival and growth of its seedlings.     

Effect of support availability,mother plant genotype and maternal support environment on the twining vine <Emphasis Type="Italic">Ipomoea purpurea</Emphasis>

In the twining vine Ipomoea purpurea we experimentally assessed the effect of support availability (a vertical stake), maternal genotype (family), and maternal environment (presence or absence of support in mother plants) on morphological traits, accounting for differences in initial seed size. While there was no effect of the maternal environment on seed size at the family level, families within each maternal environment did differ in seed size. Seeds from families of supported mothers showed higher germination than those from families of unsupported mothers. The maternal environment did not influence shoot traits but affected phenotypic plasticity in the number of leaves in response to support. Thus, whereas progeny plants from unsupported mothers did not show a response to support availability in the number of leaves, progeny from supported mother plants had a greater number of leaves once support was provided. The maternal genotype only affected the number of leaves.     

沙冬青种子萌发及幼苗生长特性

沙冬青(Ammopiptanthus mongolicus)是亚洲中部荒漠特有种,由于其分布的环境条件特殊,现存植被天然更新困难,人工育苗、造林存在一定问题。为改善种皮透水性,获得整齐一致的发芽,分别用21.5 (室温)、40、60、80和90 ℃的温水浸种5 min。结果表明: 60 ℃温水浸种效果最好,与对照相比,显著地提高了种子发芽势,并有效地降低硬实率。对比分析了15、20、25、30、35和40 ℃环境温度下种子萌发的特性,种子发芽最适温度是30 ℃,而胚根及下胚轴伸长最快的温度为25~30 ℃。较低温度下种子发芽延续时间较长,并且种子萌发后下胚轴和胚根伸长缓慢,易形成畸形(胚根严重扭曲、缢裂)幼苗。15 ℃时畸形幼苗占发芽种子数的28.1%,而30~40 ℃时仅占5.2%~8.6%。温度过高不利于种子萌发及幼苗生长,40 ℃时,吸胀种子绝大部分丧失活力,发芽种子在35 ℃以上温度下生长2~3 d,胚根胚轴组织呈水渍状坏死。另外,催芽后的种子播种在河沙中子叶出土最快,出土率可达63%,显著高于在粘质壤土中出土率(11%)。不同粒径的种子播种后幼苗生长特性表现出一定的差异,粒径5 mm以下小粒种子播种后幼苗长势弱,苗期成活率低,生长70 d后幼苗成活率仅为35.4%,而粒径5 mm以上种子同期幼苗成活率达56.2%。据上述试验结果,结合自然分布区内环境条件的资料,分析了沙冬青濒危原因,并为植被恢复育苗措施提出建议。     

Community-wide germination strategies in an alpine meadow on the eastern Qinghai-Tibet plateau: phylogenetic and life-history correlates

In this study, we built up a database of 633 species (48 families, 205 genera) from an alpine meadow on the eastern Qinghai-Tibet plateau. Our objective was to assess the effects of phylogenetic and life-history (life form, perenniality, seed size, dispersal strategy and period) background on the community-wide germination strategies. We found that the seeds of shrubs, perennials, and well-dispersed plants, and the smaller seeds germinated more and comparatively earlier. In one-way ANOVAs, phylogenetic groups explained 12% of the variance in GT (mean germination time for all seeds germinated of each species); life-history attributes, such as seed size, dispersal strategy, perenniality and life form explained 10%, 7%, 5%, and 1% respectively, and dispersal period had no significant effect on GT. Multifactorial ANOVAs revealed that the three major factors contributing to differences in GT were phylogenetic relatedness, seed size and dispersal strategy (explained 4%, 5% and 4% of the interspecific variation independently, respectively). Thus, seeds germination strategies were significantly correlated with phylogenetic and life-history relatedness. In addition, phylogenetic relatedness had close associations and interactions with seed size and dispersal strategy. Then, we think phylogeny and life-history attributes could not be considered mutual exclusively. Seed germination, like any other trait, is shaped by the natural history of the species and by the evolutionary history of the lineage. And a large percentage of the variance remained unexplained by our model, which suggested important selective factors or parameters may have been left out of this analysis. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sequential style removal in highbush blueberry,Vaccinium corymbosum L.: Effects on fertilization success and seed germination

Plants of the blueberry cultivar Bluecrop were pollinated with pollen from the cultivars Sierra and Duke under controlled conditions. After pollination, styles were removed at 6 h intervals from 1.25 days to 4.00 days. An increase in the number of developed seed was seen across time, ranging from 1 seed per fruit (s/f) at 1.25 days, to a peak of 63.4 s/f at 3.25 days in 1992, and from 0.1 s/f at 1.25 days to 56.0 s/f at 3.75 days in 1993. Germination tests of 200-seed seedlots of Bluecrop×Sierra exhibited a significant trend toward earlier germination by seed from earlier style removal times. Seed weight measured in the cross Bluecrop×Duke, however, showed decreasing weights with longer style removal times, suggesting that seed size may time of germination. A multiple regression of seed size and stylar removal time against time of germination showed that style removal time was the only significant source of variation for germination time. Results suggest that pollen vigor, as reflected in the speed of growth down the style, may have a direct bearing on characteristics of the seed and subsequent plant.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

Plant interspecies competition for sunlight: a mathematical model of canopy partitioning

We examine the influence of canopy partitioning on the outcome of competition between two plant species that interact only by mutually shading each other. This analysis is based on a Kolmogorov-type canopy partitioning model for plant species with clonal growth form and fixed vertical leaf profiles (Vance and Nevai in J. Theor. Biol., 2007, to appear). We show that canopy partitioning is necessary for the stable coexistence of the two competing plant species. We also use implicit methods to show that, under certain conditions, the species’ nullclines can intersect at most once. We use nullcline endpoint analysis to show that when the nullclines do intersect, and in such a way that they cross, then the resulting equilibrium point is always stable. We also construct surfaces that divide parameter space into regions within which the various outcomes of competition occur, and then study parameter dependence in the locations of these surfaces. The analysis presented here and in a companion paper (Nevai and Vance, The role of leaf height in plant competition for sunlight: analysis of a canopy partitioning model, in review) together shows that canopy partitioning is both necessary and, under appropriate parameter values, sufficient for the stable coexistence of two hypothetical plant species whose structure and growth are described by our model. A. L. Nevai was supported in part by the National Institutes of Health, National Research Service Award (T32-GM008185) from the National Institute of General Medical Sciences (NIGMS).     

Geographic and within-population variation in the globeflower–globeflower fly interaction: the costs and benefits of rearing pollinators’ larvae

Interspecific interactions can vary within and among populations and geographic locations, and this variation can influence the nature of the interaction (e.g. mutualistic vs. antagonistic) and its evolutionary stability. Globeflowers are exclusively pollinated by flies, whose larvae feed only on their seeds. Here we document geographic variability in costs and benefits in globeflowers in sustaining their pollinating flies throughout the range of this arctic-alpine European plant over several years. A total of 1,710 flower heads from 38 populations were analysed for their carpel, egg and seed contents. Individual and population analyses control for the confounding influences of variation in both: (1) population traits, such as fly density and egg distribution among flower heads; and (2) individuals traits, such as carpel and egg numbers per flower head. Despite considerable variation in ecological conditions and pollinator densities across populations, large proportions (range 33–58%) of seeds were released after predation, with a benefit-to-cost ratio of 3, indicating that the mutualism is stable over the whole globeflower geographical range. The stability of the mutualistic interaction relies on density-dependent competition among larvae co-developing in a flower head. This competition is revealed by a sharp decrease in the number of seeds eaten per larva with increasing larval number, and is intensified by non-uniform egg distribution among globeflowers within a population. Carpel number is highly variable across globeflowers (range 10–69), and flies lay more eggs in large flowers. Most plants within a population contribute to the rearing of pollinators, but the costs are greater for some than for others. Large globeflowers lose more seed to pollinator larvae, but also release more seed than smaller plants. The apparent alignment of interests between fly and plants (positive relationship between numbers of seed released and destroyed) is shown to hide a conflict of interest found when flower size is controlled for.     

Geographical and within-population variation in the globeflower–globeflower fly interaction: the costs and benefits of rearing pollinator’s larvae

Interspecific interactions can vary within and among populations and geographical locations, and this variation can influence the nature of the interaction (e.g. mutualistic versus antagonistic) and its evolutionary stability. Globeflowers are exclusively pollinated by flies whose larvae feed only on their seeds. Here we document geographical variability in costs and benefits in globeflowers in sustaining their pollinating flies throughout the range of this arctic-alpine European plant over several years. A total of 1,710 flower heads from 38 populations were analysed for their carpel, egg and seed contents. Individual and population analyses control for the confounding influences of variation in both: (1) population traits, such as fly density and egg distribution among flower heads; and (2) individuals traits, such as carpel and egg numbers per flower head. Despite considerable variation in ecological conditions and pollinator densities across populations, large proportions (range 33–58%) of seeds are released after predation, with a benefit-to-cost ratio of 3, indicating that the mutualism is stable over the whole globeflower geographical range. The stability of the mutualistic interaction relies on density-dependent competition among larvae co-developing in a flower head. This competition is revealed by a sharp decrease in the number of seeds eaten per larva with increasing larval number, and is intensified by non-uniform egg distribution among globeflowers within a population. Carpel number is highly variable across globeflowers (range 10–69), and flies lay more eggs in large flowers. Most plants within a population contribute to the rearing of pollinators, but some pay more than others. Large globeflowers lose more seed to pollinator larvae, but also release more seed than smaller plants. The apparent alignment of interests between fly and plant (positive relationship between numbers of seeds released and destroyed) is shown to hide a conflict of interest found when flower size is controlled for.     

Size-dependent sex allocation in a monocarpic perennial herb, <Emphasis Type="Italic">Cardiocrinum cordatum</Emphasis> (Liliaceae)

The size-dependent sex allocation model predicts that the relative resource allocation to female function often increases with plant size in animal-pollinated plants. If size effects on reproductive success vary depending on the environmental conditions, however, the size dependency may differ among populations. We tried to detect site-specific variation in size-dependent sex allocation of a monocarpic hermaphrodite with reference to light availability. Multiple flowers and fruits were sampled from the individuals of Cardiocrinum cordatum, a monocarpic understory herb, and pollen, ovule and seed production were measured with reference to the plant size in two populations. Furthermore, frequency and foraging behavior of pollinator visitation was observed. Ovule production per flower increased with plant size in both populations, while pollen production per flower increased with size only in the population under sparse canopy. Therefore, proportional allocation to male function decreased with plant size in the population under closed canopy, but did not change in the population under sparse canopy. Pollinators usually visited only one flower per plant, indicating the negligible geitonogamous pollination in this species. Although seed production under closed canopy was lower than that under sparse canopy, seed-set rate per flower and seed mass per fruit were independent of plant size in either of the populations. Size-dependent sex allocation in this species was site-specific, suggesting that not only resource storage before reproduction (i.e., plant size) but also resource availability of environment throughout the reproductive process (i.e., light availability) affect reproductive performance in this species.