Opposing natural selection from herbivores and pathogens may maintain floral-color variation in Claytonia virginica (Portulacaceae)

The maintenance of floral-color variation within natural populations is enigmatic because directional selection through pollinator preferences combined with random genetic drift should lead to the rapid loss of such variation. Fluctuating, balancing, and negative frequency-dependent selection mediated through pollinators have been identified as factors that may contribute to the maintenance of floral-color variation, and recently it has been suggested that indirect responses to selection on correlated characters through agents of selection other than pollinators may substantially shape the evolution of floral traits. Here, I provide empirical support for this latter view in Claytonia virginica (Portulacaceae) through a multiseason field study, a pollen supplementation study, and an artificial herbivory experiment. These studies show that most individuals fall into one of four discrete color classes, and suggest pollinator-mediated selection for increased floral redness in concurrent years. Floral color is also an indirect target of opposing directional selection via herbivores and pathogens that fluctuates through time. Taken together, these data suggest a novel mechanism by which floral-color variation may be maintained, and illustrate the importance of an inclusive, pluralistic view of selection when investigating the evolution of complex phenotypes.     

Cytogeography and genome size variation in the Claytonia perfoliata (Portulacaceae) polyploid complex

Background and Aims

Genome duplication is a central process in plant evolution and contributes to patterns of variation in genome size within and among lineages. Studies that combine cytogeography with genome size measurements contribute to our basic knowledge of cytotype distributions and their associations with variation in genome size.

Methods

Ploidy and genome size were assessed with direct chromosome counts and flow cytometry for 78 populations within the Claytonia perfoliata complex, comprised of three diploid taxa with numerous polyploids that range to the decaploid level. The relationship between genome size and temperature and precipitation was investigated within and across cytotypes to test for associations between environmental factors and nuclear DNA content.

Key Results

A euploid series (n = 6) of diploids to octoploids was documented through chromosome counts, and decaploids were suggested by flow cytometry. Increased variation in genome size among populations was found at higher ploidy levels, potentially associated with differential contributions of diploid parental genomes, variation in rates of genomic loss or gain, or undetected hybridization. Several accessions were detected with atypical genome sizes, including a diploid population of C. parviflora ssp. grandiflora with an 18 % smaller genome than typical, and hexaploids of C. perfoliata and C. parviflora with genomes 30 % larger than typical. There was a slight but significant association of larger genome sizes with colder winter temperature across the C. perfoliata complex as a whole, and a strong association between lower winter temperatures and large genome size for tetraploid C. parviflora.

Conclusions

The C. perfoliata complex is characterized by polyploids ranging from tetraploid to decaploid, with large magnitude variation in genome size at higher ploidy levels, associated in part with environmental variation in temperature.     

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.     

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。     

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.     

Sex-allocation trade-offs in Nigella sativa (Ranunculaceae) examined with flower manipulation experiments

Sex-allocation trade-offs have long been invoked as a primary factor underlying the evolution of separate sexes and the reduction of pollen production accompanying the evolution of selfing. In the present study, I conducted stamen and style removal experiments to explore the existence of such trade-offs in Nigella sativa, a hermaphroditic plant species whose flower structure allows early manipulation of both male and female function. Plants on which all stamens were removed at the bud stage had a higher rate of flower initiation and produced significantly heavier seeds than did plants whose flowers remained intact, apparently by using resources that were released when the stamens were removed. However, there was no effect of stamen removal on the number of flowers that reached anthesis, the total biomass allocated to seed production, or the vigour of plants in the progeny generation. In contrast, prevention of fruit production (style removal) increased the amount of biomass invested in stamen by 57% relative to plants whose flowers were allowed to set fruit. These observations verify the existence of a sexual trade-off in N. sativa but also raise the possibility that stamen-suppressing mutations sometimes lack the pleiotropic consequences of increasing female function, at least in species with large, expensive fruits.     

Trade-offs between flower number and investment to a flower in selfing and outcrossing varieties of Impatiens hypophylla (Balsaminaceae)

Floral resource allocation was compared on a whole-plant basis between two varieties of Impatiens hypophylla that differ in flower size. There were significant negative correlations between flower number and investments to a flower at both the within-population and between-variety levels. In individual flowers, var. hypophylla with larger flowers invested significantly more resources to male and pollinator-attractive functions, whereas investments to female function did not differ between the varieties. In experimental populations placed in the field, pollinators preferred the larger flowers of var. hypophylla even within the same habitat of var. microhypophylla, which has smaller flowers. There was a significant lack of observed heterozygosity only in var. microhypophylla. Thus, the outcrossing variety had more attractive but fewer flowers, while the selfing variety had less attractive but more abundant flowers.     

高寒退化草地不同海拔狼毒种群花大小与叶大小、叶数量的关系

植物花大小与叶大小、叶数量的关系反映了植物长期与环境相互作用形成的外在形态方面的适应对策,不同生境中其生长关系的变异体现了植物对异质环境的适应方式。本文研究了祁连山北坡高寒退化草地4个不同海拔狼毒(Stellera chamaejasme)种群花大小与叶大小、叶数量的关系。结果表明:随着海拔的升高,草地群落的高度、密度和地上生物量均呈先升高后降低的倒U型分布;狼毒的地上生物量、株高和叶大小均呈减小趋势,而繁殖分配、花大小和叶数量均呈增大的变化趋势;在海拔梯度上,狼毒花大小与叶数量呈极显著正相关(P0.01),与叶大小呈不显著负相关(P0.05),说明狼毒种群花大小与叶数量具有显著的依赖关系;生境对狼毒花大小与叶数量之间的依赖关系产生显著影响,高海拔的环境胁迫导致狼毒个体减小,植株通过增加花大小和叶数量、减小叶大小的资源分配策略来保证物种的繁衍。     

Smaller amphipod mothers show stronger trade-offs between offspring size and number

Trade-offs between embryo mass and number were studied in 10 populations of the freshwater amphipod Gammarus minus . Trade-offs were stronger in populations with small brooding females than in those with larger brooding females. Relationships between embryo mass and maternal body mass were also stronger in populations dominated by small versus large brooding females. These patterns are likely the result of morphological constraints, at least in part. Embryo size is more affected by brood size and maternal size in small mothers, probably because of offspring-packaging constraints associated with small brood pouches. Energy constraints appear to be less important. These results suggest that body size may not only affect the magnitude of individual life-history traits, as is well known, but also the covariance between traits.     

Flower number and floral components in ten angiosperm species: an examination of assumptions about trade-offs in reproductive evolution

A common approach to modelling reproductive evolution in flowering plants includes an implicit assumption that module number and resource allocation per module follow an inverse hyperbolic trade-off. This assumption has not been thoroughly tested. In ten herbaceous and small woody species I examined phenotypic partial correlations between flower number (measured in relation to vegetative biomass) and each of three floral components: pollen number per flower, ovule number per flower, and corolla size. Significantly negative correlations between flower number and at least one of the floral components occurred in four of the ten species. These phenotypic correlations suggest the existence of true (genetically based) trade-offs, because environmental correlations are likely to be positive, but the significant negative relationships are linear except in one case. Thus, evolutionary tradeoffs involving flower number seem likely in some cases, but there is little to indicate that hyperbolic trade-offs are common. The phenotypic patterns investigated here cannot provide definitive answers about the form of trade-offs. Nonetheless, theoretical attention to the potential evolutionary consequences of trade-offs other than the implicit hyperbolic form is needed.     

高寒退化草地狼毒种群不同海拔花大小-数量的权衡关系

权衡关系是植物生活史对策理论的基础, 花大小-数量的权衡关系对理解花的生物量分配具有重要的意义。该文利用实验生态学方法, 研究了祁连山北坡高寒退化草地4个不同海拔狼毒(Stellera chamaejasme)种群花大小与花数量间的关系。结果表明: 随着海拔升高, 草地群落的高度、密度和地上生物量均呈先升高后降低的倒U型分布, 狼毒花大小呈逐渐增大的趋势, 而花数量呈相反的变化趋势; 不同海拔样地狼毒花大小与花数量间的相关性存在差异, 海拔2700、2900和3000 m样地狼毒花大小和花数量之间均存在极显著的负相关关系(p < 0.01), 海拔2800 m样地二者之间存在显著的负相关关系(p < 0.05), 狼毒植株分配给花大小与花数量的资源间存在着“此消彼长”的权衡关系; 海拔2800 m样地是狼毒植株花大小与花数量权衡关系由低海拔的花数量多而单花生物量低向高海拔的花数量少而单花生物量高转变的区域。狼毒植株通过合理权衡花数量和花大小的资源配置, 以补偿在海拔梯度上不利因素对种群繁衍带来的影响, 从而实现种群的繁殖成功, 反映了毒杂草对异质性生境的表型可塑性机制。     

Trade-off between the biomass and number of flowers in Stellera chamaejasme along an elevation gradient in a degraded alpine grassland

权衡关系是植物生活史对策理论的基础, 花大小-数量的权衡关系对理解花的生物量分配具有重要的意义。该文利用实验生态学方法, 研究了祁连山北坡高寒退化草地4个不同海拔狼毒(Stellera chamaejasme)种群花大小与花数量间的关系。结果表明: 随着海拔升高, 草地群落的高度、密度和地上生物量均呈先升高后降低的倒U型分布, 狼毒花大小呈逐渐增大的趋势, 而花数量呈相反的变化趋势; 不同海拔样地狼毒花大小与花数量间的相关性存在差异, 海拔2700、2900和3000 m样地狼毒花大小和花数量之间均存在极显著的负相关关系(p < 0.01), 海拔2800 m样地二者之间存在显著的负相关关系(p < 0.05), 狼毒植株分配给花大小与花数量的资源间存在着“此消彼长”的权衡关系; 海拔2800 m样地是狼毒植株花大小与花数量权衡关系由低海拔的花数量多而单花生物量低向高海拔的花数量少而单花生物量高转变的区域。狼毒植株通过合理权衡花数量和花大小的资源配置, 以补偿在海拔梯度上不利因素对种群繁衍带来的影响, 从而实现种群的繁殖成功, 反映了毒杂草对异质性生境的表型可塑性机制。     

呼伦贝尔草原植物叶片大小与数量的权衡关系及其内在机制

叶片大小与数量的权衡关系是植物生长策略的基础,研究叶片大小与数量权衡关系的内在机制对于深刻理解植物生长策略具有重要意义.本研究以单个叶片干重表示叶大小,以单位茎上的叶片数量表示出叶强度,采用标准主轴回归分析方法研究呼伦贝尔草原植物叶片大小与数量的权衡关系以及内在机制.结果表明:陈旗嵯岗(典型草原)和陈旗八一(草甸草原)...     

The trade-off between number and size of offspring in humans and other primates

Life-history theory posits a fundamental trade-off between number and size of offspring that structures the variability in parental investment across and within species. We investigate this 'quantity-quality' trade-off across primates and present evidence that a similar trade-off is also found across natural-fertility human societies. Restating the classic Smith-Fretwell model in terms of allometric scaling of resource supply and offspring investment predicts an inverse scaling relation between birth rate and offspring size and a (-1/4) power scaling between birth rate and body size. We show that these theoretically predicted relationships, in particular the inverse scaling between number and size of offspring, tend to hold across increasingly finer scales of analyses (i.e. from mammals to primates to apes to humans). The advantage of this approach is that the quantity-quality trade-off in humans is placed into a general framework of parental investment that follows directly from first principles of energetic allocation.     

Genetic basis of the trade-off between offspring number and quality in the bank vole

One of the main tenets of modern life-history theory is the negative relationship (trade-off) between the number and quality of offspring produced. Theory predicts a negative genetic correlation between these traits since both are closely related to fitness of individuals. However, the genetic basis of the trade-off has only been tested to a limited extent in natural populations. We examined whether size and quality of offspring are negatively related to litter size in the bank vole Clethrionomys glareolus. First, we found a significant negative phenotypic correlation between the number and size of offspring at birth in both laboratory and field populations of the bank vole. Second, a larger size at birth decreased the maturation age of female offspring in the laboratory, and increased the probability of breeding and the size of the first litter in the field. Furthermore, manipulation of offspring size at weaning indicated that structural effects of birth size in mammals have a more profound effect on the expression of life-history traits than weaning size. Finally, in addition to the phenotypic negative correlation between the number and size of offspring, we found evidence for a negative genetic correlation between these two traits, which confirms the genetic basis of the trade-off. This negative genetic covariation may have considerable effects on the rate and direction of evolution of the two related life-historical traits.     

Effect of flower number on the pollinator attractiveness and the threshold plant size for flowering in Pertya triloba (Asteraceae)

Pertya triloba (Asteraceae) is a perennial herb growing in the understory of deciduous broad-leaved forests in central Japan. Its capitulum consists of a single floret that differs from those of most other Asteraceae species. A bagging experiment clearly showed that breeding of P. triloba required cross-pollination. By using female fertility as a measure of pollination success, the effect of an attractive floral display on the threshold plant size for flowering in P. triloba was surveyed. The number of florets per plant in a P. triloba population was artificially adjusted between one and 30. In both control plants and experimental plants with adjusted floret number, the seed set increased with an increase in the number of florets per plant and became constant (60%) when the floret number per plant was more than 10. Thus, the number of florets per plant has an important role in attracting pollinators, and more than 10 florets per plant must be the effective floral display size. In the P. triloba population, 98.5% of the flowering plants had 10 or more flowers. The effective floral display size might be determined through the mutual relationship between plants and pollinators. The number of florets per plant increased with an increase in the plant leaf area. Therefore, the threshold plant size for flowering might be determined by the productivity needed to support an attractive floral display.     

Pollinator responses to variation in floral display and flower size in dioecious Sagittaria latifolia (Alismataceae)

In animal-pollinated plants with unisexual flowers, sexual dimorphism in floral traits may be the consequence of pollinator-mediated selection. Experimental investigations of the effects of variation in flower size and floral display on pollinator visitation can provide insights into the evolution of floral dimorphism in dioecious plants. Here, we investigated pollinator responses to experimental arrays of dioecious Sagittaria latifolia in which we manipulated floral display and flower size. We also examined whether there were changes in pollinator visitation with increasing dimorphism in flower size. In S. latifolia, males have larger flowers and smaller floral displays than females. Visitation by pollinators, mainly flies and bees, was more frequent for male than for female inflorescences and increased with increasing flower size, regardless of sex. The number of insect visits per flower decreased with increasing floral display in males but remained constant in females. Greater sexual dimorphism in flower size increased visits to male inflorescences but had no influence on the number of visits to female inflorescences. These results suggest that larger flower sizes would be advantageous to both females and males, and no evidence was found that females suffer from increased flower-size dimorphism. Small daily floral displays may benefit males by allowing extended flowering periods and greater opportunities for effective pollen dispersal.