Development of polymorphic microsatellite markers in Arisaema serratum (Thunb.) Schott,Araceae

Arisaema serratum is a perennial herb capable of changing sex expression from year to year. We developed five polymorphic microsatellite markers for A. serratum to estimate male reproductive success. Variability at these loci was examined in two populations, one at Horigane and one at Kanazawa Japan; the number of alleles per locus ranged from five to 35 (the mean 21.4) in Horigane and from three to 36 (22.0) in Kanazawa. This high allelic diversity indicates that our markers are suitable for the study of male reproductive success in A. serratum.     

An Application of the Sex Allocation Theory to Arisaema serratum

Abstract The sex allocation theory based on size-specific demography is applied to sex change phenomena exhibited by Arisaema serratum , which changes sex expression according to its size. In the evolutionary analysis where adopted strategy correlates with resource level, identifying equal resource level is very important. But directly measured size does not seem to represent the plant's real resource level perfectly. In the analysis presented here, we introduce a new variable that represents the resource level. According to the sex allocation theory, we calculated the transition matrix and other demographic parameters, the stable sex-and-size distribution, and reproductive values. The obtained results were: (1) reproductive values generally increased in accordance with size, but in a different manner for different sexes; (2) in the size class where different sex expressions were adopted, reproductive values for the equal resources were the same; and (3) plant individuals seemed to adopt evolutionarily stable strategies. The adaptive nature of the plant is further discussed.     

Selective pollination by fungus gnats potentially functions as an alternative reproductive isolation among five Arisaema species

Background and AimsInterspecific difference in pollinators (pollinator isolation) is important for reproductive isolation in flowering plants. Species-specific pollination by fungus gnats has been discovered in several plant taxa, suggesting that they can contribute to reproductive isolation. Nevertheless, their contribution has not been studied in detail, partly because they are too small for field observations during flower visitation. To quantify their flower visitation, we used the genus Arisaema (Araceae) because the pitcher-like spathe of Arisaema can trap all floral visitors.MethodsWe evaluated floral visitor assemblage in an altitudinal gradient including five Arisaema species. We also examined interspecific differences in altitudinal distribution (geographic isolation) and flowering phenology (phenological isolation). To exclude the effect of interspecific differences in altitudinal distribution on floral visitor assemblage, we established ten experimental plots including the five Arisaema species in high- and low-altitude areas and collected floral visitors. We also collected floral visitors in three additional sites. Finally, we estimated the strength and contribution of these three reproductive barriers using a unified formula for reproductive isolation.Key ResultsEach Arisaema species selectively attracted different fungus gnats in the altitudinal gradient, experimental plots and additional sites. Altitudinal distribution and flowering phenology differed among the five Arisaema species, whereas the strength of geographic and phenological isolations were distinctly weaker than those in pollinator isolation. Nevertheless, the absolute contribution of pollinator isolation to total reproductive isolation was weaker than geographic and phenological isolations, because pollinator isolation functions after the two early-acting barriers in plant life history.ConclusionsOur results suggest that selective pollination by fungus gnats potentially contributes to reproductive isolation. Since geographic and phenological isolations can be disrupted by habitat disturbance and interannual climate change, the strong and stable pollinator isolation might compensate for the weakened early-acting barriers as an alternative reproductive isolation among the five Arisaema species.     

Sex Change and Population Dynamics in Arisaema (Araceae) II. An Examination on the Evolutionary Stability of Sex Changing Schedule of A. serratum (Thunb.) Schott

Abstract The evolutionary stability of the sex-changing habit of Arisaema serratum was examined. To evaluate reproductive success through male functioin properly, the mathematical formulation of Kakehashi and Harada (1987) was employed. Individuals were classified by both their size and sexual state, and the genetic contribution of individuals was estimated by calculating their reproductive value. It was shown that in the size classes where sex change actually occurs, the reproductive values of males and females are basically the same. This means that the genetic contribution of individuals in these size classes is the same whether they are male or female, and no selection pressure is working to change the male ratio in these classes. Thus, it is concluded that the size at which sex change occurs is evolutionarily stable. The adaptive significance of the size of first reproduction was also discussed.     

Temporal variation in the pollen:ovule ratios of Clarkia (Onagraceae) taxa with contrasting mating systems: field populations

Among plants, pairs of selfing vs. outcrossing sister taxa provide interesting systems in which to test predictions concerning the magnitude and direction of temporal changes in sex allocation. Although resource availability typically declines towards the end of the growing season for annual taxa, temporal changes in mating opportunities depend on mating system and should change less in selfing taxa. Consequently, given that the pollen:ovule (P:O) ratio of flowers reflects the investment in (and potential fitness pay-off due to) male vs. female function, we predicted that the P:O ratio should also be less variable among and within selfers than in closely related outcrossers. To test these predictions, we measured temporal changes in sex allocation in multiple field populations of two pairs of sister taxa in the annual flowering plant genus Clarkia (Onagraceae). In the outcrossing Clarkia unguiculata and the selfing Clarkia exilis, ovule production declined similarly from early to late buds, whereas pollen production remained constant or increased in the outcrosser but remained constant or decreased in the selfer. Consequently, the P:O ratio increased within unguiculata populations but marginally increased or stayed constant in exilis populations. In all populations of the selfing Clarkia xantiana spp. parviflora and the outcrossing C. x. spp. xantiana, both ovule and pollen production per flower declined over time. The effects of these declines on the P:O ratio, however, differed between subspecies. In each xantiana population, the mean P:O ratio did not differ between early and late flowers, although individuals varied greatly in the direction and magnitude of phenotypic change. By contrast, parviflora populations differed in the mean direction of temporal change in the P:O ratio. We found little evidence to support our initial predictions that the P:O ratio of the selfing taxa will consistently vary less than in outcrossing taxa.     

Temporal variation in phenotypic gender and expected functional gender within and among individuals in an annual plant

Background and Aims

Adaptive explanations for variation in sex allocation centre on variation in resource status and variation in the mating environment. The latter can occur when dichogamy causes siring opportunity to vary across the flowering season. In this study, it is hypothesized that the widespread tendency towards declining fruit-set from first to last flowers on plants can similarly lead to a varying mating environment by causing a temporal shift in the quality (not quantity) of siring opportunities.

Methods

A numerical model was developed to examine the effects of declining fruit-set on the expected male versus female reproductive success (functional gender) of first and last flowers on plants, and of early- and late-flowering plants. Within- and among-plant temporal variation in pollen production, ovule production and fruit-set in 70 Brassica rapa plants was then characterized to determine if trends in male and female investment mirror expected trends in functional gender.

Key Results

Under a wide range of model conditions, functional femaleness decreased sharply in the last flowers on plants, and increased from early- to late-flowering plants in the population. In B. rapa, pollen production decreased more rapidly than ovule production from first to last flowers, leading to a within-plant increase in phenotypic femaleness. Among plants, ovule production decreased from early- to late-flowering plants, causing a temporal decrease in phenotypic femaleness.

Conclusions

The numerical model confirmed that declining fruit-set can drive temporal variation in functional gender, especially among plants. The discrepancy between observed trends in phenotypic gender in B. rapa and expected functional gender predicted by the numerical model does not rule out the possibility that male reproductive success decreases with later flowering onset. If so, plants may experience selection for early flowering through male fitness.     

Current and historical factors drive variation of reproductive traits in unisexual mosses in Europe: A case study

Unisexual bryophytes provide excellent models to study the mechanisms that regulate the frequency of sexual versusasexual reproduction in plants, and their ecological and evolutionary implications. Here, we determined sex expression, phenotypic sex ratio, and individual shoot traits in 242 populations of the cosmopolitan moss Pseudoscleropodium purum spanning its whole distributional range. We tested whether niche differentiation, sex-specific differences in shoot size, and biogeographical history explained the spatial variation of reproductive traits. We observed high levels of sex expression and predominantly female-biased populations, although both traits showed high intraspecific variation among populations. Sex expression and sex ratio were partly explained by current macroscale environmental variation, with male shoots being less frequent at the higher end of the environmental gradients defined by the current distribution of the species. Female bias in population sex ratio was significantly lower in areas recolonized after the last glacial maximum (recent populations) than in glacial refugia (long-term persistent populations). We demonstrated that reproductive trait variation in perennial unisexual mosses is partially driven by macroscale and historical environmental variation. Based on our results, we hypothesize that sexual dimorphism in environmental tolerance and vegetative growth contribute to sex ratio bias over time, constraining the chances of sexual reproduction, especially in long-term persistent populations. Further studies combining genetic analyses and population monitoring should improve our understanding of the implications of the intraspecific variation in the frequency of sexual versusasexual reproduction in bryophyte population fitness and eco-evolutionary dynamics.     

Inter-genotype variation in reproductive response to crowding among Daphnia pulex

Crowding is known to have a major influence on reproduction in the freshwater microcrustacean Daphnia pulex. We analyzed reproductive output of six different D. pulex genotypes under two different density regimes in the laboratory. Four of these genotypes reproduce via obligate parthenogenesis, allowing thorough analysis of the life history strategies of some asexual lines. Among 30,109 neonate offspring and 1041 resting egg ephippia collected, several trends were evident. Crowding induced increased resting egg production and reduced neonate offspring production among all genotypes. Offspring sex ratios grew more male-biased with maternal age. The extent, but not direction, of each of these trends varied among genotypes. Offspring sex ratios, and the very direction in which they changed in response to crowding, differed significantly among genotypes with some genotypes producing more and others fewer males in response to crowding. Obligately parthenogenetic genotypes seemed to respond to the crowding stimulus in similar ways as the facultatively parthenogenetic genotypes, as expected from the sexual origins of their genomes. The inter-genotype variation in life-history traits observed in this and other investigations calls into question the common practice of extrapolating results from a single Daphnia genotype to an entire species. Our findings are considered in the context of other research in the field of environmental influences on Daphnia reproduction with a review of representative literature.     

Variation in structural gender in the hermaphrodite Helleborus foetidus (Ranunculaceae): within- and among-population patterns

In hermaphrodite plants, variations in structural gender (defined as the ratio between male and female gametes) may occur at different levels (among flowers, plants, and populations). In this study, we investigated variation in four traits influencing structural gender (number of carpels, ovules per carpel, stamens, and pollen grains per stamen) within and among six distant populations of the hermaphrodite perennial herb Helleborus foetidus (Ranunculaceae) in the Iberian Peninsula. Our results show that the four traits investigated varied significantly at all levels considered. Traits influencing the female sexual component (number of carpels and ovules per carpel) showed greater variation at the lowermost levels (within flower and plant) than traits influencing the male component, which in turn varied more markedly among populations. Number of carpels per flower and number of pollen grains per anther were the most important traits affecting between-plant variation in structural gender. There was no evidence of significant plant-level trade-offs or correlations between the various male and female traits, which covaried differently across populations. The observed between-population variation in structural gender of Helleborus foetidus can be explained as a consequence of differences in self-pollination levels related to a flowers mating environment.     

苔藓植物生殖生态学研究

近年来苔藓植物生殖生态学研究主要集中于繁育系统、生殖代价与对策,以及不同生殖方式对种群遗传变异的影响等方面。生殖结构的原始性及其对水分的独特需求,以及雌雄异株比例较高等导致苔藓植物中有性生殖比例偏低;雌配子体很少完成整个有性生殖过程,其“真实的生殖代价”主要指雌性性表达(雌配子发生)的能耗,并且显著低于雄性性表达;基于对资源有效分配的生殖对策而导致雌性偏向及部分孢子体败育。无性生殖有利于不同生境条件下有效种群的发展与维持,其多样化的繁殖方式导致复杂的种群动态。苔藓植物具有较高的种群遗传多样性,生殖方式与种群遗传变异无直接因果关系,孢子与无性繁殖体不同的散布能力对于种群间遗传分化具有一定的影响。     

Heritable variation of sex ratio in a polychaete worm

Ophryotrocha labronica is a gonochoristic polychaete worm whose sex determining mechanism and sex ratio control are supposed to be polygenic. From a lab population, whose sex ratio (i.e., proportion of males) was 0.5, the estimate of sex ratio heritability by offspring-father regression was 0.54 ± 0.15 and by offspring-mother regression was not significantly different from 0. Estimate of sex ratio repeatability between successive broods of a pair was 0.64 ± 0.33. Since female parents do not contribute in any way to the variability of sex ratio, sex ratio variation seems to be largely a paternal character. On the basis of these estimates we advance the hypothesis that in this species sex is determined by a multilocus genetic system, allowing the combined effects of a female major sex gene (which could give rise to a form of female heterogamety) and masculinizing modifiers. The hypothesis that the male sex has the least canalised sexual differentiation is supported by the observation that some old males developed oocytes.     

Linking the evolution of gender variation to floral development

BACKGROUND AND AIMS: In the present review, I have endeavoured to conduct a joint assessment of the thinking underlying the evolutionary genetics of gender polymorphism and the developmental genetics of gender determination. It is my hope, through highlighting the historical development of ideas in two related but somewhat disparate sets of scientific literature, to encourage a synthetic perspective that integrates the two. SCOPE: An overview is provided of various theories on the evolution of sex polymorphism and examples of evidence that has been brought to bear in support of them. Current knowledge on floral development is summarized, with an emphasis on gender variation. Finally, an attempt is made to integrate the two perspectives with the hope that it will encourage future research at the interface. CONCLUSIONS: Evolutionary models of gender evolution have, of necessity, posited genetic effects that are relatively simple in their impacts. Emerging insights from developmental genetics have demonstrated that the underlying reality is a more complex matrix of interacting factors. The study of gender variation in plants is poised for significant advance through the integration of these two perspectives. Bringing genomic tools to bear on population-level processes, we may finally develop a comprehensive perspective on the evolution of floral gender.     

Preliminary data on the influence of rearing temperature on the growth and reproductive status of fathead minnows Pimephales promelas

An investigation into the influence of temperature on the growth and reproductive status of the fathead minnow Pimephales promelas revealed that, while there was no clear effect of treatment on sex differentiation, ovarian tissue from female fish reared under the highest temperature regime contained large amounts of undefined tissue containing no germ cells. Furthermore, both male and female fish exhibited differences in length mass, condition and somatic indices, and in the expression of secondary sexual characteristics. The patterns observed are discussed in the context of climate change.     

Genetic variation and reproductive patterns in wetland mosses suggest efficient initial colonization of disturbed sites

To understand colonization processes, it is critical to fully assess the role of dispersal in shaping biogeographical patterns at the gene, individual, population, and community levels. We test two alternative hypotheses (H I and H II) for the colonization of disturbed sites by clonal plants, by analyzing intraspecific genetic variation in one and reproductive traits in two typical fen mosses with separate sexes and intermittent spore dispersal, comparing disturbed, early‐succession (limed) fens and late‐successional rich fens. H I suggests initial colonization of disturbed sites by diverse genotypes of which fewer remain in late‐successional fens and an initially balanced sex ratio that develops into a possibly skewed population sex ratio. H II suggests initial colonization by few genotypes and gradual accumulation of additional genotypes and an initially skewed sex ratio that alters into the species‐specific sex ratio, during succession. Under both scenarios, we expect enhanced sexual reproduction in late‐successional fens due to resource gains and decreased intermate distances when clones expand. We show that the intraspecific genetic diversity, assessed by two molecular markers, in Scorpidium cossonii was higher and the genetic variation among sites was smaller in disturbed than late‐successional rich fens. Sex ratio was balanced in S. cossonii and Campylium stellatum in disturbed fens and skewed in C. stellatum in late‐successional fens, thus supporting H I. In line with our prediction, sex expression incidence was higher in, and sporophytes were confined to, late‐succession compared to disturbed rich fens. Late‐successional S. cossonii sites had more within‐site patches with two or more genotypes, and both species displayed higher sex expression levels in late‐successional than in disturbed sites. We conclude that diverse genotypes and both sexes disperse efficiently to, and successfully colonize new sites, while patterns of genetic variation and sexual reproduction in late‐successional rich fens are gradually shaped by local conditions and interactions over extended time periods.     

两性具有不同出生率和死亡率的种群动态

以往的种群动力学模型均隐含假设性比为1：1,而实际上并非总如此,不同性别的出生和死亡是不完全相同的。文中就考虑了两性具有不同出生率和死亡率的种群动态问题。可以知道种群动态只是受雌性控制,与雄性无关。     

Ecological and phylogenetic determinants of life-history patterns among ten loricariid species

We analysed the influence of ecological factors, phylogenetic history and trade-offs between traits on the life-history variation among 10 loricariid species of the middle Paraná River. We measured eight life-history variables and classified the life-history strategies following the equilibrium–periodic–opportunistic (EPO) model. Principal-component analysis of life-history traits segregated species along a gradient from small opportunistic (low fecundity, low parental investment) to large equilibrium (low-medium fecundity, high parental investment) species. A clear periodic strategist was absent in the analysed assemblage. Variation partitioning by canonical phylogenetic ordination analysis showed both a component of variation uniquely explained by phylogenetic history (PH; 32.2%) and a component shared between PH and ecological factors (EF; 37%). The EPO model is a useful tool for predicting correlations among life-history traits and understanding potential demographic responses of species to environmental variation. Life-history patterns observed throughout Loricariidae suggests that this family has diversified across all three endpoint strategies of the EPO model. Our study indicates that evolutionary lineage affiliation at the level of subfamily can be a strong predictor of the life-history strategy used by each species.     

Sex-ratio variation and reproductive costs in relation to density in a forest-dwelling population of red deer (Cervus elaphus)

For dimorphic species in which the variance in reproductivesuccess of males is more pronounced than that of females, theoriesof adaptive variation in sex ratio predict that mothers shouldinvest more heavily in sons than in daughters. By using harvestdata from a forest-dwelling red deer population that experienceda marked reduction in population density we tested the hypothesisthat adaptive sex-ratio variation should occur only when populationsare much below carrying capacity. More specifically, we testedwhether at low density, females in better than average conditionwere more likely to produce male offspring and to invest inindividual sons rather than were females in poorer than averagecondition. We also investigated female reproductive costs arisingfrom a decrease either in body mass or in reproduction. We didnot find any support for a biased sex ratio or investment towardmale calves by high-quality mothers at any population density.Costs of reproduction in terms of body mass and pregnancy rateswere only detectable for females that reproduced as yearlingsand not for those that reproduced as adults. Our results thereforedo not support the hypothesis of adaptive sex-ratio variationin a population living below carrying capacity. The four-folddifference in party sizes (defined as the number of deer aggregatein the same party in which no individual was more than 50 mfrom any other) observed in our population living in a closedforest habitat compared with populations living in more openhabitats previously studied might account for such a discrepancy.We suggest that a smaller party size may decrease the intensityof sexual selection and could be the proximal cause for thelack of adaptive sex-ratio variation we report for the populationstudied here.     

Effects of diet and host access on fecundity and lifespan in two fruit fly species with different life‐history patterns

The reproductive ability of female tephritids can be limited and prevented by denying access to host plants and restricting the dietary precursors of vitellogenesis. The mechanisms underlying the delayed egg production in each case are initiated by different physiological processes that are anticipated to have dissimilar effects on lifespan and reproductive ability later in life. The egg‐laying abilities of laboratory‐reared females of the Mediterranean fruit fly (Ceratitis capitata Wiedmann) and melon fly (Bactrocera cucurbitae Coquillett) from Hawaii are delayed or suppressed by limiting access to host fruits and dietary protein. In each case, this is expected to prevent the loss of lifespan associated with reproduction until protein or hosts are introduced. Two trends are observed in each species: first, access to protein at eclosion leads to a greater probability of survival and a higher reproductive ability than if it is delayed and, second, delayed host access reduces lifetime reproductive ability without improving life expectancy. When host access and protein availability are delayed, the rate of reproductive senescence is reduced in the medfly, whereas the rate of reproductive senescence is generally increased in the melon fly. Overall, delaying reproduction lowers the fitness of females by constraining their fecundity for the remainder of the lifespan without extending the lifespan. © 2013 The Royal Entomological Society