Long-term variation in seed mass and seed production in populations of Paris quadrifolia

Seed production of the perennial herb Paris quadrifolia L. (Liliaceae) was investigated in five populations in northern Poland. The long-term seed production per square metre differed significantly among populations and years. Moreover, throughout the 7-year study period, 30% of both whole ripe fruits and seeds alone were predated. Variation in seed mass per fruit in both space and time was significant. Throughout the 7-year study, nearly all the marked individuals produced fruits once every 2 years. The most frequent break between fruiting was 2 years and the longest was 5 years. Only the mean seed mass in fruits of the same individual varied significantly over subsequent years. In the five populations, the number of ovules, number of seeds in the fruit and seed mass varied significantly between populations. However seed/ovule ratio did not differ in fruits in the five populations. The seed mass/number trade-off in fruits was strongly partially correlated when the effect of total seed mass was considered. Breeding experiments suggest that P. quadrifolia has a substantial capacity for both inbreeding and outbreeding. There were no significant differences in the seed/ovule ratio, seed number or seed mass in fruits produced from bagged or control flowers. However, both seed/ovule ratio and number of seeds were significantly lower in fruits from emasculated flowers.     

Fruit to flower ratios and trade-offs in size and number

Summary Hermaphroditic flowering plants commonly produce many more flowers than fruits. To interpret this observation, I suggest an explanation based on trade-offs occurring between the size and the number of flowers and fruits that mature when resources are limiting. I make this suggestion concrete with the aid of a phenotypic model analysed using the ESS approach. The model includes resource allocations to male and female reproductive structures at the time of flowering and to fruit maturation. The formulation allows for non-linear relations between opportunities for fertility gain on the one hand and flower and fruit size and number on the other. Results of the model indicate that fewer fruits will be matured than flowers produced, even in the absence of previously suggested factors such as pollinator limitation and bet-hedging in the face of environmental variability. Model results emphasize the distinction between flower versus inflorescence contributions to male and female reproductive success. Analysis also shows that the fruit to flower ratio in hermaphroditic plants will be lower than that in female plants, in accord with broad taxonomic surveys. Finally, results obtained lead to empirically testable predictions about the relationship between flower size, fruit size and the fruit to flower ratio. Methods to test these predictions are discussed.     

天马自然保护区常见秋熟植物种子大小变异分析

以种子千粒质量为指标,对安徽天马自然保护区61种常见秋熟植物(种子数量在500粒以上)的种子大小进行了测定,并从种子大小与生活型、分类群和种子产量的相关性等方面研究了种子大小的变异规律.结果显示:61种植物的种子大小存在很大变异,种子千粒质量的最小值和最大值分别为0.015和79.370 g;根据种子千粒质量可将种子大小划分成A(0～0.05 g)、B(0.05 ～0.5 g)、C(0.5 ～5 g)、D(5 ～50 g)和E(50 ～500 g)5个等级,分别占总种数的8.20％、31.15％、36.06％、21.31％和3.28％,呈正态分布.61种植物包含乔木、藤本、灌木和草本4种生活型,种子千粒质量平均值分别为24.753、6.884、6.812和1.601 g,不同生活型间的种子大小存在极显著差异.61种植物分属33科,不同科间的种子大小具有极显著差异,但各科间的种子大小与分类系统发育方向没有显著相关性.种子较小且种子产量较高的种类有茵陈蒿(Artemisia capillaris Thunb.)、奇蒿(A.anomala S.Moore)、薄雪火绒草(Leontopodium japonicum Miq.)、华东藨草(Scirpus karuizawensis Makino)和庐山藨草(S.lushanensis Ohwi)等,显示种子大小与种子产量呈显著的负相关.     

Game theory provides no explanation for seed size variation in grasslands

Game theoretical models have been suggested to explain the maintenance of a remarkable variation in seed size across species in most types of vegetation. According to these models, which are based on the existence of a trade-off between seed size and seed number; smaller-seeded species can invade any species mixture due to their numerical advantage, and larger-seeded species can invade any species mixture due to their competitive superiority over smaller-seeded species during recruitment. However, till now, there is very limited evidence for seed size effects on recruitment interactions among different species. An experiment was designed using 16 species in Swedish grasslands, varying 384-fold in seed size. Species were sown pairwise and alone in disturbed versus undisturbed small plots in grazed versus ungrazed grassland, and the resulting recruitment was recorded. Seedling densities in the experimental plots were within the natural range. Both disturbance and grazing had a positive effect on recruitment. Seed size did not affect recruitment except from a tendency that increasing seed size made recruitment less dependent on disturbance. Recruitment of smaller-seeded species was not affected by the presence of larger-seeded species. Larger-seeded species did not generally win in direct contest. These results suggest that game theoretical models do not explain maintenance of seed size variation across species in these grasslands. Alternative explanations for seed size variation are that either small-scale heterogeneity provide conditions favouring a range of different seed sizes or other attributes than seed size effectively determine recruitment.     

Co-evolution of seed size and seed predation

Using the evolutionarily stable strategy (ESS) approach in a model for the co-evolution of seed size and seed predation, I show that seed size variation within individual plants is favoured if there is a trade-off in the predator's attack rate for different seed sizes. A single seed size is not evolutionarily stable because a predator that is optimally adapted to one particular seed size cannot prevent invasion by plants with a different seed size. The model generates the following predictions. The ESS consists of a continuous range of seed sizes. Small seeds tend to be attacked more frequently than big seeds. Plants with many resources and plants with low (frequency-independent) juvenile mortality have more variable seeds than plants with few resources and a high juvenile mortality. Seed size variation is higher in fluctuating populations regulated by seed predation alone than in stable populations (partially) regulated by seedling competition. Predator searching behaviour does not directly affect the ESS seed size range, but may have an indirect effect by affecting population stability or the significance of seedling competition as a population regulating mechanism. Moreover, seed size distributions are found to be more skewed in favour of small seeds if predation is spatially non-uniform than if predation is more even. Application of the model to systems of several co-evolving plant and predator species is discussed.     

Hamilton's rule, imprinting and parent-offspring conflict over seed mass in partially selfing plants

When genes in the offspring control the provisioning of the seed, the optimal seed size can be calculated exactly by applying Hamilton's rule. When seed size is a compromise between mother and offspring, we predict that outcrossing plant species produce larger seeds than selfers. This trend was found in the British flora and in a number of well-studied plant families. The analysis was extended to imprinting, a conditional strategy in which a gene in the offspring takes more resources when derived from the father than from the mother. The conditions for imprinting to be selected were rather restrictive. The analysis is relevant for the current debate about the evolution of imprinting in Arabidopsis thaliana.     

Pollen and seed dispersal among dispersed plants

The ecological significance of spacing among plants in contributing to the maintenance of species richness, particularly in tropical forests, has received considerable attention that has largely focussed on distance- and density-dependent seed and seedling mortality. More recently it has become apparent that plant spacing is also relevant to pollination, which often constrains seed production. While seed and seedling survival is reduced at high conspecific densities, pollination success, by contrast, is positively correlated to local conspecific density. Distance-dependent mechanisms acting on pollination and seed production have now been described for a variety of plants, with relatively isolated plants or fragmented populations generally suffering reduced fecundity due to pollen limitation. Yet there is considerable variability in the vulnerability of plant species to pollination failure, which may be a function of breeding system, life history, the pollination vector, the degree of specialisation among plants and their pollinators, and other indirect effects of habitat change acting on plants or pollinators. As reduced tree densities and population fragmentation are common outcomes of anthropogenically altered landscapes, understanding how pollination processes are affected in such degraded landscapes can inform effective conservation and management of remaining natural areas.     

Paternal effects in Arabidopsis indicate that offspring can influence their own size

The existence of genetic variation in offspring size in plants and animals is puzzling because offspring size is often strongly associated with fitness and expected to be under stabilizing selection. An explanation for variation in seed size is conflict between parents and between parents and offspring. However, for this hypothesis to be true, it must be shown that the offspring genotype can affect its own size. The existence of paternal effects would support this hypothesis, but these have rarely been shown. Using a diallel cross among four natural accessions of Arabidopsis thaliana we show that maternal, paternal and positional effects jointly influence seed size, number and the frequency of seed abortion. We found that seed abortion (%) depends on the combination of maternal and paternal genotypes, suggesting the existence of mate choice or epistatic incompatibility among accessions of A. thaliana. In addition, since paternal genotype explains approximately 10 per cent of the variation in seed size, we propose that A. thaliana''s offspring must influence the amount of resources allocated to themselves. Identification of paternal effects in Arabidopsis should facilitate dissection of the genetic mechanisms involved in paternal effects.     

A model for nectar secretion in animal-pollinated plants

Summary A game theoretic model was developed for nectar secretion in animal-pollinated plants in order to examine how the total amount of resources allocated to flowers affects the spread of nectarless plants. It was assumed that pollinators concentrate on patches whose nectar rewards are relatively large compared to other patches and if pollinators visit a patch, they concentrate on the plants whose nectar rewards are relatively large compared to other plants in the patch. It was shown that plants are more likely to secrete nectar in populations where the total amount of resources allocated to flowers is large. It was also shown that strong interplant competition, strong interpatch competition and the nectar discrimination of the pollinators are also important factors for nectar secretion. However, if the total amount of resources allocated to flowers is sufficiently large, plants would secrete nectar even if competition is not very strong and nectar discrimination is not so precise.     

Hidden responses to environmental variation: maternal effects reveal species niche dimensions

Species responses to fluctuating environments structure population and community dynamics in variable ecosystems. Although offspring number is commonly used to measure these responses, maternal effects on offspring quality may be an important but largely unrecognised determinant of long‐term population growth. We selected 29 species across a Mediterranean annual plant phylogeny, and grew populations of each species in wet and dry conditions to determine responses in seed number and maternal effects (seed size, seed dormancy, and seedling growth). Maternal effects were evident in over 40% of species, but only 24% responded through seed number. Despite a strong trade‐off between seed size and seed number among species, there was no consistent trade‐off within species; we observed correlations that ranged from positive to negative. Overall, species in this plant guild show a complex range of responses to environmental variation that may be underestimated when only seed number responses are considered.     

壳斗科三种植物种子大小对昆虫寄生及种子存活率的影响

种子内的寄生昆虫可以严重影响种子的发育、损害种子活力。种子足余策略理论认为大种子有利于抵御和适应昆虫寄生取食,但动物最优觅食理论推测,大种子更易遭受昆虫寄生。为对这两种对立观点进行验证,本实验以青冈、苦槠和麻栎各2个种群的种子为材料,对昆虫寄生与完好种子间的体积和萌发率进行比较,并对寄生种子萌发率与种子体积的关系进行了分析。结果显示:(1)在6个种群的种子中,只有松阳麻栎和青冈种群的寄生种子体积大于完好种子,其余4个种群的寄生种子体积小于完好种子,但这种差异不显著;(2)所有寄生种子的整体萌发率(18%)显著低于完好种子(45.66%)(P<0.001),在不同种群内,寄生种子的萌发率也分别显著低于完好种子。(3)比较同种植物体积差异显著的寄生种子的萌发率发现,大种子总比小种子具有更高的萌发率,但差异不显著;在不同植物的寄生种子间比较时,体积最大的麻栎种子萌发率显著高于体积较小的青冈和苦槠种子。研究结果表明,象虫在种子上产卵时对大种子没有选择偏好,在昆虫寄生取食严重损害种子活力的压力下,大种子比小种子具有更强的耐受力。     

Geographic variation in reproductive traits and trade-offs between size and number of eggs of the Chinese cobra (Naja atra)

We collected gravid Chinese cobras (Naja atra) from one island (Dinghai) and three mainland (Yiwu, Lishui and Quanzhou) populations in south‐eastern China to study geographical variation in female reproductive traits and the trade‐off between the size and number of eggs. We then conducted an common experiment on cobras from two of the four populations to further identify factors contributing to the observed trade‐offs. The mean size (snout–vent length) of the smallest five reproductive females increased with increasing latitude. Oviposition occurred between late June and early August, with females from the warmer localities laying eggs earlier than those from the colder localities. Maternal size was a major determinant of the reproductive investment in all populations, with larger females producing not only more but also larger eggs. Clutch size was more variable than egg size within and among populations. The observed geographical variation in clutch size, egg size, clutch mass and post‐oviposition body condition was not a simple consequence of variation in maternal size among populations, because interpopulation differences in these traits were still evident when the influence of maternal size was removed. The upper limit to reproductive investment was more likely to be set by the space availability in the island population, but by the resource availability in the three mainland populations. Trade‐offs between size and number of eggs were detected in all populations, with females that had larger clutches for their size having smaller eggs. Egg size at any given level of relative fecundity differed among populations, primarily because of interpopulation differences in the resource availability rather than the space availability. Except for the timing date of oviposition and the mean size of the smallest five reproductive females, all other examined traits did not vary in a geographically continuous trend. The common garden experiment, which standardized environmental factors, synchronized the timing date of oviposition, but it did not modify the conclusion drawn from the gravid females collected from the field. The observed geographical variation in the female reproductive traits could be attributed to the consequence of the effects of either proximate or ultimate factors. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 27–40.     

水蓼花大小在花序和个体上的变化及其与数量权衡关系研究

探讨一年生植物水蓼花大小在花序上和个体上的变化及花大小与花数量的权衡关系。在54株植物个体上随机选取一花序,在花序的基部、中部和顶部各选取1～2朵花,花大小(生物量)以基部最大(0.851mg),顶部最小(0.715mg),可能是由结构效应引起的。在每个体上随机采集4～20朵花,以其均值表示植物个体的花大小,花大小不随植物个体大小变化而变化。花朵展示和总花数均随个体增大而增加。在控制植物个体大小(地上部分营养器官生物量)后,没有发现花朵展示或花数量与花大小之间的权衡关系,表明个体资源水平的差异可能掩盖了花数量与大小间的权衡关系。     

A comparative study of seed yield parameters in Arabidopsis thaliana mutants and transgenics

Because seed yield is the major factor determining the commercial success of grain crop cultivars, there is a large interest to obtain more understanding of the genetic factors underlying this trait. Despite many studies, mainly in the model plant Arabidopsis thaliana, have reported transgenes and mutants with effects on seed number and/or seed size, knowledge about seed yield parameters remains fragmented. This study investigated the effect of 46 genes, either in gain- and/or loss-of-function situations, with a total of 64 Arabidopsis lines being examined for seed phenotypes such as seed size, seed number per silique, number of inflorescences, number of branches on the main inflorescence and number of siliques. Sixteen of the 46 genes, examined in 14 Arabidopsis lines, were reported earlier to directly affect in seed size and/or seed number or to indirectly affect seed yield by their involvement in biomass production. Other genes involved in vegetative growth, flower or inflorescence development or cell division were hypothesized to potentially affect the final seed size and seed number. Analysis of this comprehensive data set shows that of the 14 lines previously described to be affected in seed size or seed number, only nine showed a comparable effect. Overall, this study provides the community with a useful resource for identifying genes with effects on seed yield and candidate genes underlying seed QTL. In addition, this study highlights the need for more thorough analysis of genes affecting seed yield.