The relative importance of architecture and resource competition in allocation to pollen and ovule number within inflorescences of Hosta ventricosa varies with the resource pools

Background and Aims

Allocation of resources to floral traits often declines distally within inflorescences in flowering plants. Architecture and resource competition have been proposed as underlying mechanisms. The aim of the present study is to assess the relative importance of resource competition and architectural effects in pollen and ovule production on racemes of Hosta ventricosa, an apomictic perennial herb.

Methods

Combinations of two defoliation treatments (intact and defoliated) and two fruit-set treatments (no-fruit and fruit) were created, and the roles of architecture and resource competition at each resource level were assessed.

Key Results

Pollen and ovule number per flower increased after defoliation, but pollen to ovule ratio per flower did not change. Pollen, ovules and the pollen to ovule ratio per flower declined distally on racemes at each resource level. In the intact treatment, fruit development of early flowers did not affect either pollen or ovule number of late flowers. In the defoliated treatment, fruit development of early flowers reduced both pollen and ovule numbers of late flowers due to over-compensation caused by defoliation. Late flowers on defoliated fruit racemes produced less pollen than intact fruit racemes but the same number of ovules; therefore, the reduction in pollen number was not caused by over-compensation. In addition, the fruit-set rate of early flowers during flowering was higher in intact racemes than in defoliated racemes.

Conclusions

In flowering plants, the relative importance of architecture and resource competition in allocation to pollen and ovules may vary with the resource pools or the overall resource availability of maternal plants.     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因.为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并...     

蒙古沙冬青花序内性分配的变化、传粉者运动与繁殖成功

性分配理论主要研究繁殖资源在雌雄功能间的最优分配,从雌雄功能的角度考虑其个体适合度.对花序内不同部位花的雌性与雄性资源分配变化的研究,对于我们理解植物采取哪种繁殖对策保障繁殖成功具有重要意义.本文对生长在中国科学院吐鲁番沙漠植物园内的蒙古沙冬青(Ammopiptanthus mongolicus)连续开花花序内不同部位...     

Paternal reproductive success drives sex allocation in a wild mammal

Parents should bias sex allocation toward offspring of the sex most likely to provide higher fitness returns. Trivers and Willard proposed that for polygynous mammals, females should adjust sex‐ratio at conception or bias allocation of resources toward the most profitable sex, according to their own body condition. However, the possibility that mammalian fathers may influence sex allocation has seldom been considered. Here, we show that the probability of having a son increased from 0.31 to 0.60 with sire reproductive success in wild bighorn sheep (Ovis canadensis). Furthermore, our results suggest that females fertilized by relatively unsuccessful sires allocated more energy during lactation to daughters than to sons, while the opposite occurred for females fertilized by successful sires. The pattern of sex‐biased offspring production appears adaptive because paternal reproductive success reduced the fitness of daughters and increased the average annual weaning success of sons, independently of maternal allocation to the offspring. Our results illustrate that sex allocation can be driven by paternal phenotype, with profound influences on the strength of sexual selection and on conflicts of interest between parents.     

Temperature-related sex allocation shifts in a recovering keystone species,Pinus palustris

Summary

In the light of all that has been discovered about the mechanism of evolution it has become tempting to follow Darwin's lead and to “see no limit to this power“. Yet a careful examination of situations in which evolution is known to be occurring in some species, shows complete absence of evolution in others. This not because these species have not had the opportunity; in many situations there may even be uncolonised bare space.

The explanation must lie in the supply of appropriate variation. A tacit assumption of evolution by natural selection is that the necessary variation is always available. Yet there is no a priori justification for this. Evidence from populations in nature, particularly of species which are potential colonists of old metal mine workings and similar metal contaminated habitats, shows that the species that successfully colonise these habitats, by the evolution of metal tolerant populations, possess within their normal populations the necessary variation. But those species which fail to colonise these habitats, despite the opportunity, do not possess this variation. This applies also at the level of the population, in the replicated evolutionary situations occurring under electricity pylons.

Such evidence, together with arguments from theory, suggests that the failures of evolution have been as important as its successes in moulding the living world as we see it today, and that the reasons for failure must be sought at the molecular level in limitations to the origin of new variation.     

Reproductive ecology of the endangered Alpine species Eryngium alpinum L. (Apiaceae): phenology, gene dispersal and reproductive success

BACKGROUND AND AIMS: Eryngium alpinum (Apiaceae) is an endangered perennial, characteristic of the Alpine flora. Because the breeding system influences both demographic (reproductive success) and genetic (inbreeding depression, evolutionary potential) parameters that are crucial for population maintenance, the reproductive ecology of E. alpinum was investigated. Specifically, the aims of the study were (1) to determine the factors (resources and/or pollen) limiting plant fitness; and (2) to assess the potential for gene flow within a plant, within a patch of plants, and across a whole valley where the species is abundant. METHODS: Field experiments were performed at two sites in the Fournel valley, France, over three consecutive years. Studies included a phenological survey, observations of pollinators (visitation rates and flight distances), dispersal of a fluorescent powder used as a pollen analogue, the use of seed traps, determination of the pollen/ovule ratio, and an experiment to test whether seed production is limited by pollen and/or by resources. KEY RESULTS: E. alpinum is pollinated by generalist pollinators, visitation rates are very high and seed set is resource- rather than pollen-limited. The short flights of honeybees indicate a high potential for geitonogamy, and low pollen and seed dispersals suggest strong genetic structure over short distances. These results are interpreted in the light of previous molecular markers studies, which, in contrast, showed complete outcrossing and high genetic homogeneity. CONCLUSION:S. The study highlights the usefulness of adopting several complementary approaches to understanding the dynamic processes at work in natural populations, and the conservation implications for E. alpinum are emphasized. Although the studied populations do not seem threatened in the near future, long-term monitoring appears necessary to assess the impact of habitat fragmentation. Moreover, this study provides useful baseline data for future investigations in smaller and more isolated populations.     

Florally rich habitats reduce insect pollination and the reproductive success of isolated plants

Landscape heterogeneity in floral communities has the potential to modify pollinator behavior. Pollinator foraging varies with the diversity, abundance, and spatial configuration of floral resources. However, the implications of this variation for pollen transfer and ultimately the reproductive success of insect pollinated plants remains unclear, especially for species which are rare or isolated in the landscape. We used a landscape‐scale experiment, coupled with microsatellite genotyping, to explore how the floral richness of habitats affected pollinator behavior and pollination effectiveness. Small arrays of the partially self‐compatible plant Californian poppy (Eschscholzia californica) were introduced across a landscape gradient to simulate rare, spatially isolated populations. The effects on pollinator activity, outcrossing, and plant reproduction were measured. In florally rich habitats, we found reduced pollen movement between plants, leading to fewer long‐distance pollination events, lower plant outcrossing, and a higher incidence of pollen limitation. This pattern indicates a potential reduction in per capita pollinator visitation, as suggested by the lower activity densities and richness of pollinators observed within florally rich habitats. In addition, seed production reduced by a factor of 1.8 in plants within florally rich habitats and progeny germination reduced by a factor of 1.2. We show this to be a consequence of self‐fertilization within the partially self‐compatible plant, E. californica. These findings indicate that locally rare plants are at a competitive disadvantage within florally rich habitats because neighboring plant species disrupt conspecific mating by co‐opting pollinators. Ultimately, this Allee effect may play an important role in determining the long‐term persistence of rarer plants in the landscape, both in terms of seed production and viability. Community context therefore requires consideration when designing and implementing conservation management for plants which are comparatively rare in the landscape.     

Patterns and causes of spatial variation in the reproductive success of a desert annual

Summary This study examines patterns and causes of variation in the reproductive success of the desert annual Stipa capensis. Three nested scales of variation were analyzed: variation between individuals of the same plot, variation between different plots of the same habitat, and variation between different habitats in the same region. Perturbation experiments (irrigation and neighbors removal) were performed to test the effects of heterogeneity in soil water and neighborhood competition on the magnitude of variation in each scale. The results demonstrate that variation of reproductive success was highest within plots, lowest between plots, and moderate between habitats. Soil water heterogeneity contributed to spatial variation in all scales but was most important for differences between habitats. Neighborhood competition increased the variation within plots, but decreased the variation between habitats. The results further demonstrate that water limitation was negatively correlated with the position of the habitat along the run-off/run-on gradient. An opposite trend was obtained for the effect of competition.     

Female reproductive success and the evolution of mating-type frequencies in tristylous populations

In tristylous populations, mating-type frequencies are governed by negative frequency-dependent selection typically resulting in equal morph ratios at equilibrium. However, Narcissus triandrus generally exhibits long-styled (L)-biased populations with a deficiency of the mid-styled (M)-morph. Here we used a pollen-transfer model and measurements of female fertility in natural populations to investigate whether these uneven morph ratios were associated with variation in female reproductive success. Our theoretical analysis demonstrated that morph ratio bias can result from maternal fitness differences among the morphs, and that these effects were magnified by asymmetrical mating. In nine out of 15 populations of N. triandrus, seed set differed significantly among the morphs, but pollen limitation occurred in only two of 11 populations investigated. Average seed set of the M-morph was positively associated with its frequency in populations. Flower size was negatively correlated with the seed set of the M-morph. Our results suggest that interactions between mating patterns and female fertility are responsible for variation in morph frequencies and loss of the M-morph from tristylous populations of N. triandrus.     

Seasonal variation in the sex allocation of a neotropical solitary bee

We carried out a field study on the life history and sex allocationof the ground-nesting solitary bee Diadasina distincta (Hymenoptera: Anthophoridae).This species is multivoltine, undergoing five generations a yearbetween February and September. The numerical sex ratio of thisspecies was female biased overall (approximately 38% males)and showed a strong and consistent seasonal pattern. The numericalsex ratio was extremely female biased (approximately 20% males)from February until May, and then slightly male biased (approximately60% males) from June until September. Females were 3.26 timesthe size of males, and so the overall investment ratio was female biasedthroughout the year. The overall female bias and seasonal variationin sex allocation is unlikely to be explained by models thatinvoke overlapping generations or competition between brothersfor mates (local mate competition). We suggest that a possibleexplanation for the female bias in the early part of the seasonis local resource enhancement (LRE): nesting near larger numbersof sisters reduces parasitism. LRE is likely to decrease in importancein the later part of the season, when the biased numerical and investmentratios may be explained by models in which male and female offspringgain different fitness returns from resources invested.     

Floral density,pollen limitation,and reproductive success in<Emphasis Type="Italic"> Trillium grandiflorum</Emphasis>

Decreases in floral density can disrupt mutualistic interactions between plants and their pollinators, and decrease reproductive success. I addressed the relationship between floral density and plant reproductive success using two experimental approaches: a pollen supplementation experiment in 12 populations of Trillium grandiflorum that naturally varied in floral density, and a transplant experiment in which floral density was manipulated in plots at four experimental sites. In the pollen supplementation experiments, the degree of pollen limitation, in terms of fruit set and seed set, decreased with floral density. Further, in the experimental sites, plant reproductive success increased asymptotically with floral density. These results demonstrate the value of simultaneously conducting experiments in both experimental sites and natural populations to understand how population density influences plant reproductive success. Factors that reduce the density of this perennial herb, such as habitat fragmentation and herbivory by white-tailed deer (Odocoileus virginianus), should be expected to limit its reproduction.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Morph-ratio variation, population size and female reproductive success in distylous Pulmonaria officinalis (Boraginaceae)

Theory predicts that morph ratios in heterostylous populations are governed by negative frequency-dependent selection typically resulting in equal morph ratios at equilibrium. Previous work on the distylous perennial herb Pulmonaria officinalis, however, showed asymmetric mating between floral morphs and a weak self-incompatibility system, with the long-styled morph (L-morph) producing significantly higher seed set following intramorph crosses and even selfing than the short-styled morph (S-morph), two aspects thought to affect female fecundity and morph-ratio variation. Here, we evaluated morph ratios and population size of all known P. officinalis populations in the northern part of Belgium. Morph ratios deviated significantly from 1:1 (range 0.09-1 L-morph frequency, mean = 0.58). Relative fecundity of the S-morph (i.e. mean seed set of the S-morph/mean seed set of the L-morph) was on average 0.73, was positively related to the frequency of the L-morph, and reached 1 (similar levels of female fecundity) at an average L-morph frequency of 0.66 in the population. As some small populations had the S-morph in majority, our results suggest that local morph ratios are influenced both by the relative fecundity of L- and S-morph individuals and by stochastic processes in small populations.     

Genetic variation in sex allocation in a parasitic wasp: variation in sex pattern within sequences of oviposition

In order to maximize their fitness under Local Mate Competition (LMC), arrhenotokous female wasps have to produce a precise sex ratio when encountering hosts. Recent progress in the theory of hymenopterous parasitoid reproduction suggest that they manage to do it by laying male and female eggs in a particular order and that such reproductive strategies are adaptive. Therefore, the determinism of such sequential patterns would be regulated by genetic control on which natural selection could act. To test this hypothesis, sequences of oviposition were recorded in a set ofTrichogramma brassicae Bezdenko (Hymenoptera; Trichogrammatidae) females and in their daughters by providing themEphestia kuehniella Zeller (Lepidoptera; Pyralidae) eggs. In order to describe accurately sex pattern within these oviposition sequences, I present a joined non-parametric and multivariate statistical method. It is shown thatT. brassicae females do not produce male and female eggs in random sequences. Moreover, the way they organize the sequence of the sexes in their progeny seems to be under a strong genetic control. The evolutionary consequences of such results are discussed.     

The effects of inbreeding, genetic dissimilarity and phenotype on male reproductive success in a dioecious plant

Pollen fate can strongly affect the genetic structure of populations with restricted gene flow and significant inbreeding risk. We established an experimental population of inbred and outbred Silene latifolia plants to evaluate the effects of (i) inbreeding depression, (ii) phenotypic variation and (iii) relatedness between mates on male fitness under natural pollination. Paternity analysis revealed that outbred males sired significantly more offspring than inbred males. Independently of the effects of inbreeding, male fitness depended on several male traits, including a sexually dimorphic (flower number) and a gametophytic trait (in vitro pollen germination rate). In addition, full-sib matings were less frequent than randomly expected. Thus, inbreeding, phenotype and genetic dissimilarity simultaneously affect male fitness in this animal-pollinated plant. While inbreeding depression might threaten population persistence, the deficiency of effective matings between sibs and the higher fitness of outbred males will reduce its occurrence and counter genetic erosion.     

Determinants of reproductive success and offspring sex in a turtle with environmental sex determination

Despite the importance of maternal effects in evolution, and knowledge of links among nest site choice, timing of nesting, offspring sex, and reproductive success in animals with environmental sex determination, these attributes have not been rigorously studied in a combined and natural context. To address this need we studied the relationships between three maternal traits (nest site choice, lay date, and nest depth) and two fitness‐related attributes of offspring (hatchling sex and embryonic survival) in the riverine turtle Carettochelys insculpta, a species with temperature‐dependent sex determination, for four years. Predation and flooding were the major sources of embryonic mortality in 191 nests. Embryonic survival was influenced by both lay date and nest site choice: in one year when nesting began later than average, nests laid later and at lower elevations were destroyed by early wet season river rises. In other years early nesting precluded flood mortality. However, turtles did not nest at the highest available elevations, and a field experiment confirmed that turtles were constrained to nest at lower elevations where they could construct a nest chamber. The principal determinant of hatchling sex in 140 nests was lay date, which in turn was apparently related to the magnitude of the previous wet season(s). Clutches laid earlier in the season (a female's first clutch) produced mainly males, while later clutches (her second clutch) yielded mostly females, due to seasonal increases in air temperatures. Accordingly, later nesting produced female‐biased hatchling sex ratios in 1996, while earlier nesting resulted in sex ratios near unity in the other years. However, all‐female nests were more likely to be flooded than mixed‐sex or all‐male nests in years when nesting was late. In conclusion, we found evidence that the position of two maternal trait distributions (elevation of the nest site and lay date), associated with the reproductive strategy of C. insculpta, reflect a combination of natural selection, physical constraints, and phenotypic plasticity. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 1–16.     

Variation in sex ratio, morph-specific reproductive ecology and an experimental test of frequency-dependence in the gynodioecious Kallstroemia grandiflora (Zygophyllaceae)

An enduring puzzle in gynodioecious species is the great variation in female frequency seen among populations. We quantified sex ratio in 44 populations of gynodioecious Kallstroemia grandiflora. Then, we measured pollinator visitation, pollen deposition, autonomous selfing rate and pollen limitation of females. Finally, using experimental populations, we tested whether female fitness responds to the frequency of female plants. We found broad variability in sex ratio among populations (0-44% female). Hermaphrodite flowers received more pollinator visits and pollen grains than females, and bagged hermaphrodite flowers produced fruits. However, we found no evidence of pollen limitation in females. In experimental populations, female plants showed no evidence of frequency-dependent pollinator visitation, fruit set, seed set or total seed mass. These results do not support frequency-dependent variation in fitness as a major mechanism affecting female frequencies in K. grandiflora. Within the context of this study, pollinators are abundant and pollinator movement appears to operate at a large enough scale to overcome the potential reproductive disadvantages of producing solely female flowers.     

Genotypic variation for reproductive characters,and the influence of pollen-ovule ratio on selfing rate in rape seed (Brassica napus)

The variance component for genotypic effects and covariable plant vigor were estimated for pollen production, number of ovules, the weight of the flower without anther sacs and pistil, and the gender of the whole plant, in the rape seed variety Topas. There was a significant effect of the genotype for all measured characters. We did not find any trade-off between the male and female functions. Small plants were relatively more male-biased than large plants. The weight of the flower without anther sacs or pistil was positively correlated with pollen production. The selfing rates of the lines were positively correlated with the pollen-ovule ratio, suggesting that the selfing rate of individual plants is positively correlated with pollen-ovule ratio, thus supporting the “mass-action” model. It is proposed that the observed increase in the population selfing rate during the flowering season for some species, including rape seed, can be caused by an overall decrease in male investment.     

Culled males,infant mortality and reproductive success in a pre-industrial Finnish population

Theoretical and empirical literature asserts that the sex ratio (i.e. M/F) at birth gauges the strength of selection in utero and cohort quality of males that survive to birth. We report the first individual-level test in humans, using detailed life-history data, of the ‘culled cohort’ hypothesis that males born to low annual sex ratio cohorts show lower than expected infant mortality and greater than expected lifetime reproductive success. We applied time-series and structural equation methods to a unique multigenerational dataset of a natural fertility population in nineteenth century Finland. We find that, consistent with culled cohorts, a 1 s.d. decline in the annual cohort sex ratio precedes an 8% decrease in the risk of male infant mortality. Males born to lower cohort sex ratios also successfully raised 4% more offspring to reproductive age than did males born to higher cohort sex ratios. The offspring result, however, falls just outside conventional levels of statistical significance. In historical Finland, the cohort sex ratio gauges selection against males in utero and predicts male infant mortality. The reproductive success findings, however, provide weak support for an evolutionarily adaptive explanation of male culling in utero.