THE EFFECT OF SEED SIZE AND MATERNAL SOURCE ON INDIVIDUAL SIZE IN A POPULATION OF LUDWIGIA LEPTOCARPA (ONAGRACEAE)

Seed size is normally distributed for many annual species, while mature plant size is frequently positively skewed. A study was conducted to determine the influence of seed size and the role of genetic differences in determining relative seedling size for Ludwigia leptocarpa. Seed size had a significant effect on percentage germination and time of seed germination but no effect on dry weight or leaf area of seedlings. Seed size and spacing had a significant effect on seedling dry weight for plants grown under competition, while relative day of emergence had no effect. Familial (genetic) differences were found in average seed weight between maternal plants, but not in average number of days to germination, average weight of seeds which germinated, or shoot dry weight. It is concluded that neither seed size alone nor genetic differences between plants are directly responsible for the development of size hierarchies in Ludwigia leptocarpa populations. Large seed size does convey an advantage in growth when plants from seeds of differing initial size interact.     

Seed size as an indicator of seed quality: a case study of Primula veris

Seed size is a widely accepted measure of seed quality, because many earlier studies have shown that large seeds have high seedling survival, growth and establishment. We tested whether ovule loss increases size of the remaining seeds and whether such size increase affects seedling establishment. We removed all except one flower from inflorescences of Primula veris L. (Primulaceae), a perennial hemicryptophyte herb, at a late stage of flowering. Flower removal (FR) increased seed size by 33% compared to the control plants. We then divided the seeds within each treatment to small, middle-sized and large seeds and carried out a sowing experiment in the field. Within each experimental group, seedling establishment was positively associated with seed size. However, despite size differences, seeds from the FR and control groups had the same seedling establishment probability. Seeds from FR plants had a higher seedling emergence in May than those from control plants, but the number of seedlings alive per sowing plot in the late summer was the same in both experimental groups. Increase in seed mass after partial FR thus did not enhance seedling performance, although seed size variation due to other causes was positively correlated with seedling establishment. Further studies are needed to show whether plastic changes of seed size are usually adaptive or not.     

Seed size as an indicator of seed quality: a case study of Primula veris

Seed size is a widely accepted measure of seed quality, because many earlier studies have shown that large seeds have high seedling survival, growth and establishment. We tested whether ovule loss increases size of the remaining seeds and whether such size increase affects seedling establishment. We removed all except one flower from inflorescences of Primula veris L. (Primulaceae), a perennial hemicryptophyte herb, at a late stage of flowering. Flower removal (FR) increased seed size by 33% compared to the control plants. We then divided the seeds within each treatment to small, middle-sized and large seeds and carried out a sowing experiment in the field. Within each experimental group, seedling establishment was positively associated with seed size. However, despite size differences, seeds from the FR and control groups had the same seedling establishment probability. Seeds from FR plants had a higher seedling emergence in May than those from control plants, but the number of seedlings alive per sowing plot in the late summer was the same in both experimental groups. Increase in seed mass after partial FR thus did not enhance seedling performance, although seed size variation due to other causes was positively correlated with seedling establishment. Further studies are needed to show whether plastic changes of seed size are usually adaptive or not.     

Seed number and environmental conditions do not explain seed size variability for the invasive herb Lupinus polyphyllus

Intraspecific variation in seed size may result from life-history constraints or environmental conditions experienced. This variation in seed size is likely to affect the early stage of invasion as seed size may contribute to the success or failure of population establishment. However, only a few studies have examined seed size variability and its causes and consequences for invaders so far. Using the invasive herb Lupinus polyphyllus, we estimated seed mass variation within and among 39 populations from two different geographic regions in a part of the invaded range. We empirically and experimentally evaluated the effect of seed number and environmental conditions (e.g. geographic region, habitat type, intraspecific competition) on seed mass, emergence and seedling performance. Seed mass varied threefold, being largest among individual plants within populations and smallest among populations. Variation in seed mass was neither related to seed number nor the environmental conditions examined, but led to differences in offspring performance, with emergence and seedling size increasing with seed mass. Larger L. polyphyllus seeds were better establishers than smaller seeds regardless of environmental conditions, indicating that the success of L. polyphyllus invasions is likely to depend positively on seed mass. Our results suggest that some plant species such as the invasive L. polyphyllus may not show an adaptive response in seed mass to resources or environmental conditions, which may partly explain their ability to colonise a range of different habitats.     

QUANTITATIVE GENETICS OF SEQUENTIAL LIFE-HISTORY AND JUVENILE TRAITS IN THE PARTIALLY SELFING PERENNIAL,AQUILEGIA CAERULEA

We determined the genetic basis of several traits related to overall fitness of Aquilegia caerulea, a perennial herb of the Rocky Mountains in western North America. To obtain measures of heritability relevant to the evolutionary potential of wild populations, we performed full and partial diallel crosses and studied progeny performance in the field. Based on a joint analysis of two designs with a total of 18 parents and 102 crosses, we detected significant maternal variance for seed mass and emergence time, but this component was negligible for later-expressed traits. Low heritability and evidence that maternal effects on seed mass are largely environmental suggest that in this population there is little evolutionary potential for change in seed mass under conditions experienced during the study. Seed mass varied depending on particular combinations of parents and cross direction. Such an interaction can have several different biological interpretations, including that particular maternal parents selectively provision embryos sired by particular pollen genotypes. Width of the first true leaf after 4 wk of growth and leaf size of juvenile plants at years one and two were significantly heritable and positively genetically correlated. Juvenile survival exhibited significant dominance variance, as expected from evidence of inbreeding depression in this trait. In contrast, for other traits that exhibit inbreeding depression in this population (seed mass and third-year leaf size), dominance variance was negligible.     

Phenotypic plasticity and population differentiation in seeds and seedlings of the grass Anthoxanthum odoratum

Summary Seeds of Anthoxanthum odoratum were transplanted reciprocally between a xeric and a mesic field population that were genetically differentiated in adult traits. In one experiment seeds were reciprocally buried in bags in the soil, in a second experiment seeds were reciprocally sown in small plots. For most traits, site effects were much larger than seed-source effects. Germination, emergence, mortality of buried seed and recruitment were significantly higher at the xeric site than at the mesic site, irrespective of population of origin. Seed dormancy, was significantly higher for seed originating from the mesic than from the xeric population. Seedling recruits originating from the xeric population tended to be larger at both sites. Fecundity of seedling recruits depended on the environment; fecundities of plants growing in the xeric site had more than double the fecundity of plants growing in the mesic site. Phenotypic plasticity rather than population differences determined variation in performance in the seed and seedling stages.     

Mediation of seed provisioning in the transmission of environmental maternal effects in Maritime pine (Pinus pinaster Aiton)

Although maternal environmental effects are increasingly recognized as an important source of phenotypic variation with relevant impacts in evolutionary processes, their relevance in long-lived plants such as pine trees is largely unknown. Here, we used a powerful sample size and a strong quantitative genetic approach to analyse the sources of variation of early seedling performance and to identify seed mass (SM)-dependent and -independent maternal environmental effects in Maritime pine. We measured SM of 8924 individual seeds collected from 10 genotypes clonally replicated in two environments of contrasting quality (favourable and stressful), and we measured seedling growth rate and biomass allocation to roots and shoots. SM was extremely variable (up to 14-fold) and strongly determined by the maternal environment and the genotype of the mother tree. The favourable maternal environment led to larger cones, larger seeds and reduced SM variability. The maternal environment also determined the offspring phenotype, with seedlings coming from the favourable environment being 35% larger and with greater root/shoot ratio. Transgenerational plasticity appears, thus, to be a relevant source of phenotypic variation in the early performance of this pine species. Seed provisioning explained most of the effect of the maternal environment on seedling total biomass. Environmental maternal effects on seedling biomass allocation were, however, determined through SM-independent mechanisms, suggesting that other epigenetic regulation channels may be involved.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

Time – size tradeoffs: a phylogenetic comparative study of flowering time, plant height and seed mass in a north-temperate flora

Parents face a timing problem as to when they should begin devoting resources from their own growth and survival to mating and offspring development. Seed mass and number, as well as maternal survival via plant size, are dependent on time for development. The time available in the favorable season will also affect the size of the developing juveniles and their survival through the unfavorable season. Flowering time may thus represent the outcome of such a time partitioning problem. We analyzed correlations between flowering onset time, seed mass, and plant height in a north-temperate flora, using both cross-species comparisons and phylogenetic comparative methods. Among perennial herbs, flowering onset time was negatively correlated with seed mass (i.e. plants with larger seeds started flowering earlier) while flowering onset time was positively correlated with plant height. Neither of these correlations was found among woody plants. Among annual plants, flowering onset time was positively correlated with seed mass. Cross-species and phylogenetically informed analyses largely agreed, except that flowering onset time was also positively correlated with plant height among annuals in the cross-species analysis. The different signs of the correlations between flowering onset time and seed mass (compar. gee regression coefficient=−7.8) and flowering onset time and plant height (compar. gee regression coefficient=+30.5) for perennial herbs, indicate that the duration of the growth season may underlie a tradeoff between maternal size and offspring size in perennial herbs, and we discuss how the partitioning of the season between parents and offspring may explain the association between early flowering and larger seed mass among these plants.     

Post‐dispersal seed predation and its relations with seed traits: a thirty‐species‐comparative study

Post‐dispersal seed predation is a key process determining the variability in seed survival in forests, where most seeds are handled by rodents. Seed predation is thought to affect seedling regeneration, colonization ability and spatial distribution of plants. Basic seed traits are the essential factors affecting rodent foraging preferences and thus seed survival and seedling recruitment. Many studies have discussed several seed traits and their effects upon seed predation by rodents. However, the results of those previous studies are usually equivocal, likely because few seed traits and/or plant species tend to be incorporated into these studies. In order to elucidate the relationships between seed predation and seed traits, we surveyed the predation of 48 600 seeds in a natural pine forest, belonging to 30 species, for three consecutive years. The results demonstrated that: (i) seed size and seed coat hardness did not significantly affect seed predation; (ii) total phenolics had a negative effect upon seed predation; (iii) positive effects of nitrogen content upon seed predation were found. From our study, it seems that the better strategy to prevent heavy predation is for plants to produce seeds with higher total phenolics content rather than physical defenses (i.e. hard seed coat) or larger seeds. Additionally, rodent foraging preference may depend more on Nitrogen content than other nutrient content of seeds.     

Quantitative genetics of seed size variation inPhlox

Summary Because seed size is often associated with survival and reproduction in plant populations, genetic variation for seed size may be reduced or eliminated by natural selection. To test this hypothesis we assessed genetic sources of variation in seed size in a population ofPhlox drummondii to determine whether genetic differences among seeds influence the size they attain. A diallel cross among 12 plants from a population at Bastrop, Texas, USA allowed us to partition variance in the mass of seeds among several genetic and parental effects. We found no evidence of additive genetic variance or dominance genetic variance for seed mass in the contribution of plants to their offspring. Extranuclear maternal effects accounted for 56% of the variance in seed mass. A small interaction was observed between seed genotype and maternal plant. Results of this study support theory that predicts little genetic variation for traits associated with fitness.     

Intrinsic and extrinsic drivers of recruitment across the distribution range of a seed-dimorphic herb

Recruitment of new individuals, through germination and seedling survival, is a key process for short-lived plants. Here, we analyzed intraspecific variation in recruitment across the latitudinal range of Plantago coronopus, a widespread herb that produces both large basal seeds with a mucilaginous coat and small apical seeds without coat. We experimentally tested the effects of seed traits and water availability on recruitment, by using seeds from a wide environmental stress gradient from N Africa to N Europe. Our experiments were carried out in controlled environmental conditions and in dunes where the species naturally occurs. Water shortage decreased seed germination and seedling survival for all populations, showing the importance of water supply for P. coronopus. Basal seeds showed higher and faster germination rates than apical seeds. Since among-population variation in seed mass was not related to potential germination rate, it is the mucilaginous coat rather than size difference that likely drives the differential success between seed morphs. Seed mass positively affected seedling survival instead, but only in controlled conditions with regular water supply. An experiment in a dune showed indeed that the highest survival corresponded to the local population and not the one with the largest seeds. Our results demonstrate that both intrinsic and extrinsic factors drive inter-population variation in the early life stages of this short-lived plant, allowing it to adapt across the environmentally heterogeneous distribution range. Gathering information on intraspecific variation in recruitment-related traits will help us to understand and predict plant responses in a context of climatic change.     

EFFECTS OF PARENTAGE AND SIZE OF THE POLLEN LOAD ON PROGENY PERFORMANCE IN CAMPANULA AMERICANA

To examine the effects of maternal and paternal parentage and the size of the pollen load on seed size and weight and on progeny performance we conducted a controlled crossing experiment using a natural population of Campanula americana. We found that seed size was positively correlated with early seedling performance for all but one of traits we measured (days to emergence), but was not significantly correlated with any of the later vegetative measures or reproductive output. We detected significant effects due to the maternal parent for the vegetative traits days to emergence, days to first leaf, and final plant height, as well as total seed weight, and mean seed weight per fruit. Significant paternal effects were found for all of the seedling traits except number of leaves after vernalization. The progeny from fruits receiving high pollen loads significantly outperformed the progeny from fruits receiving low pollen loads for the traits days to first and second leaf, numbers of leaves after vernalization, and days to first flower. These results not only demonstrate the importance of parentage and seed weight on progeny performance, but also indicate that variations in the size of the pollen load may be important in seedling establishment in natural populations.     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

Effects of seed traits variation on seedling performance of the invasive weed,Ambrosia artemisiifolia L.

Seedling performance can determine the survival of a juvenile plant and impact adult plant performance. Understanding the factors that may impact seedling performance is thus critical, especially for annuals, opportunists or invasive plant species. Seedling performance can vary among mothers or populations in response to environmental conditions or under the influence of seed traits. However, very few studies have investigated seed traits variations and their consequences on seedling performance. Specifically, the following questions have been addressed by this work: 1) How the seed traits of the invasive Ambrosia artemisiifolia L. vary among mothers and populations, as well as along the latitude; 2) How do seed traits influence seedling performance; 3) Is the influence on seedlings temperature dependent. With seeds from nine Western Europe ruderal populations, seed traits that can influence seedling development were measured. The seeds were sown into growth chambers with warmer or colder temperature treatments. During seedling growth, performance-related traits were measured. A high variability in seed traits was highlighted. Variation was determined by the mother identity and population, but not latitude. Together, the temperature, population and the identity of the mother had an effect on seedling performance. Seed traits had a relative impact on seedling performance, but this did not appear to be temperature dependent. Seedling performance exhibited a strong plastic response to the temperature, was shaped by the identity of the mother and the population, and was influenced by a number of seed traits.     

Adaptive traits of wild barley plants of Mediterranean and desert origin

Reciprocal introduction of seeds and seedlings was used to test for local adaptation and to identify a set of co-adapted traits of Mediterranean and desert ecotypes of wild barley Hordeum spontaneum. Evidence for local adaptation was found in seedling introductions into intact environments and from ecotype colonization success in the first generation after seed dispersal. Estimates of fitness were obtained at particular stages of the life cycle (seed, seedling and adult). Experiments that manipulated the environment (vegetation removal, different plant density) demonstrated the intensity and direction of natural selection in different life history episodes, but there was no strong evidence for local adaptation under these circumstances. The observed genetically determined differences between Mediterranean and desert ecotypes can be summarized as the following: reproductive output was higher in desert plants, with smaller seeds than in Mediterranean plants. There was a higher competitive ability of Mediterranean than desert plants. Plants of desert origin had significant reductions in yield when grown in mixed stands with Mediterranean plants; no such effect was observed for plants of Mediterranean origin. Seed germination and seedling survival was lower in seeds of desert origin. This was due to both: genetically determined higher dormancy of desert seeds and a trade-off between no. of seeds and their size (directly related to seed/seedling vigour).     

Sabal palmetto seed size: causes of variation,choices of predators,and consequences for seedlings

High variation in seed size, as is common among angiosperms, may be maintained in a plant species when several factors select for seed size. Variation may also result from differences among adult plants, such as nutrient and water availability or the amount of photosynthetic tissue. In a study of Sabal palmetto seed ecology I found high seed size variation both within- and among-palms, and investigated possible factors maintaining this variation. Seed size was positively correlated with the number of leaves on parent palms. Larger seeds produced more vigorous seedlings that had greater leaf length, area, and mass, and greater root mass. Caryobruchus gleditsiae (Bruchidae: Coleoptera), whose larvae develop within palm seeds, preferentially oviposited on larger seeds, which in turn produced larger beetle offspring. By choosing the largest seeds available, ovipositing beetles thus affect both the quantity and the quality of seeds available for recruitment. I conclude that because beetle predation selects against large seeds, while larger seeds promote seedling vigor, the maintenance of seed size variation may be an adaptation of S. palmetto promoting both seed escape from predators and seedling vigor.     

Effect of seed position in spikelet on life history of Eremopyrum distans (Poaceae) from the cold desert of north-west China

Background and Aims

Most studies on seed position-dependent effects have focused on germination characteristics. Our aim was to determine the effects of seed position in the spikelet on differences in timing of germination and on the ecological life history of the grass Eremopyrum distans in its cold desert habitat.

Methods

For seeds in three spikelet positions, morphology, mass and dormancy/germination characteristics were determined in the laboratory, and seeds planted in field plots with and without watering were followed to reproduction to investigate seedling emergence and survival, plant size and seed production.

Key Results

After maturation, of the seeds within the spikelet, basal ones (group 1) are the largest and have the highest proportion with physiological dormancy, while distal ones (group 3) are the smallest and have the highest proportion of non-dormant seeds. A higher percentage of seeds after-ripened in groups 2 and 3 than in group 1. Seeds sown in the field in early summer and watered at short, regular intervals germinated primarily in autumn, while those under natural soil moisture conditions germinated only in spring. Both cohorts completed their life cycle in early summer. Seeds in group 1 had lower percentages of seedling emergence and higher percentages of seedling survival than those in groups 2 and 3. Also, plants from group 1 seeds were larger and produced more seeds per plant than those from groups 2 and 3.

Conclusions

Seed position-dependent mass was associated with quantitative differences in several life history traits of E. distans. The environmentally enforced (low soil moisture) delay of germination from autumn to spring results in a reduction in fitness via reduction in number of seeds produced per plant.     

Are the best dispersers the best colonizers? Seed mass,dispersal and establishment in <Emphasis Type="Italic">Carduus</Emphasis> thistles

Negative correlations between dispersal and establishment are often reported in the plant literature; smaller seeds tend to disperse better but germinate less well, and produce smaller seedlings. However, because dispersal capacity is often quantified using proxies, such as the settling velocity of wind-dispersed seeds, little is known about the exact shape of this negative relationship, and how it is modified by other plant traits and environmental conditions. We studied the dispersal-establishment relationship in two wind-dispersed thistles (Carduus nutans and Carduus acanthoides). We applied a mechanistic wind dispersal model (WALD) to seeds released under a range of environmental conditions, and tested germination and seedling growth under standardized conditions in a greenhouse. Dispersal distance and establishment (germination and seedling growth) were not significantly correlated, although in both species smaller seeds dispersed farther, and showed lower germination and lower seedling growth rates. This apparent paradox can partly be explained by the significant influence of other factors such as release height and environment (wind and vegetation), which explained more variation in dispersal than did terminal velocity. Another potential explanation is the variation in seed traits: germination is strongly positively related to seed mass, weakly positively related to plume loading, but not significantly related to terminal velocity. This weakening of the correlation with germination is due to additional layers of trait (co)variability: for instance, seed mass and pappus size are positively correlated, and thus big seeds partially compensate for the negative effect of seed mass with larger pappi. Our mechanistic approach can thus lead to a better understanding of both potentially opposing selection pressures on traits like seed mass, and diluting effects of other seed, plant and environmental factors.     

Seed mass and emergence time effects on performance of Panicum dichotomiflorum Michx. across environments

Summary The effects of seed mass and emergence time on growth and reproduction of the annual, C₄ grass, Panicum dichotomiflorum Michx., were investigated in an experimental field study. Individual seeds of P. dichotomiflorum of known mass were sown into marked plots in four newly-abandoned fields that differed in soil nutrient availability, soil moisture and the time of the year in which they were abandoned. Seedling emergence was monitored daily for four weeks following sowing. In none of these fields was there any relationship between seed mass and emergence time. Analyses of covariance indicated that the effects of seed mass and emergence date on plant performance varied among fields. Seed mass affected performance in one field (SOUTH EARLY) where abiotic stress after emergence was high. In contrast, later emergence significantly reduced performance in all four fields. We used a path analysis to decompose the direct and indirect effects of seed mass and emergence time on plant performance in each field. The path analysis revealed that in all four fields, final weight had a significant positive effect on seed production. The effect of seed mass and emergence time had variable (and often opposite effects) on these traits. Seed mass only had a significant positive, direct effect on seed production in the most abiotically stressful field (SOUTH LATE). In the other fields, the effects of seed mass on fecundity were mediated by the effect on final weight. Thus, the effects of seed mass and emergence time on growth and reproduction in P. dichotomiflorum vary across environments.