Density-dependent changes in sex ratio in Colias lesbia (Lepidoptera: Pieridae)

Abstract.

• 1 The changes in sex ratio of a population of Colias lesbia were studied for four consecutive years.
• 2 A constant sex ratio was found in the samples taken in the 34 days following the last harvest of the alfalfa.
• 3 For the samples taken 35 or more days after the harvest the sex ratio decreases as the total number of adults caught increases.
• 4 The indirect evidence collected suggests that differences in emigration tendencies between sexes play a major role in determining the sex ratio in this species.

     

The involvement of phytohormones in the plant sex regulation

The data concerning the plant sex regulation by phytohormones are presented. Functioning of signaling pathways regulating floral development and sex expression, including those with phytohormone involvement, are considered. The role of phytohormones in the functioning of systems and mechanisms of sex regulation is analyzed. The examples of sex reversion by plant treatment with phytohormones are presented. It is demonstrated that many genes determining sex encode proteins involved in the phytohormone metabolism. The significance of phytohormone investigation for the understanding of molecular mechanisms of plant sex regulation is discussed.     

Density-dependent impact of the human malaria parasite Plasmodium falciparum gametocyte sex ratio on mosquito infection rates

Malaria parasites produce male and female life cycle stages (gametocytes) that must fertilize to achieve successful colonization of the mosquito. Gametocyte sex ratios have been shown to be under strong selection pressure both as an adaptive response to a worsening blood environment for transmission and according to the number of co-infecting clones in the vertebrate. Evidence for an impact of sex ratio on the transmission success of Plasmodium falciparum has, however, been more controversial. Theoretical models of fertilization predict that increasingly male sex ratios will be favoured at low gametocyte densities to ensure fertilization. Here, we analyse in vitro transmission studies of P. falciparum to Anopheles gambiae mosquitoes and test this prediction. We find that there is a discernible effect of sex ratio on transmission but which is dependent upon the gametocyte density. While increasingly male sex ratios do give higher transmission success at low gametocyte densities, they reduce success at higher densities. This therefore provides empirical confirmation that sex ratio has an immediate impact on transmission success and that it is density-dependent. Identifying the signals used by the parasite to alter its sex ratio is essential to determine the success of transmission-blocking vaccines that aim to impede the fertilization process.     

Kin recognition in an annual plant

Kin recognition is important in animal social systems. However, though plants often compete with kin, there has been as yet no direct evidence that plants recognize kin in competitive interactions. Here we show in the annual plant Cakile edentula, allocation to roots increased when groups of strangers shared a common pot, but not when groups of siblings shared a pot. Our results demonstrate that plants can discriminate kin in competitive interactions and indicate that the root interactions may provide the cue for kin recognition. Because greater root allocation is argued to increase below-ground competitive ability, the results are consistent with kin selection.     

Soil moisture and sex ratio in a plant with nuclear-cytoplasmic sex inheritance

I investigated whether soil moisture affects relative fitness of females and hermaphrodites and sex ratio in a gynodioecious plant with nuclear-cytoplasmic sex inheritance. I contrast these results with those from species with strictly nuclear sex inheritance. I performed a manipulative watering experiment on seed fitness of the two sexes, and field studies measuring seed fitness and sex ratio as a function of soil moisture. In the dry site, watered hermaphrodites produced approximately twice as many seeds as unwatered hermaphrodites, with little treatment effect on female seed production. Over a natural soil moisture gradient, the ratio of female to hermaphrodite seed production was higher in dry than in wet sites. These data show that the seed fitness advantage of females is a function of soil moisture. Despite this, regression of soil moisture on the sex ratio of 23 populations was not significant. These results indicate a sex-dependent effect of soil moisture on resource allocation to seeds that does not translate into a strong effect on sex ratio. This is consistent with theory based on genomic conflict in which sex ratios are predicted to be only partly determined by fitness differences of the sexes.     

Survivorship patterns in an annual plant community

Summary Age-specific survivorship was studied in a number of plant species which occur in a distinct, predominantly annual plant community of Texas, in association with the activities of pocket gophers. Of eleven species analyzed in detail, all functioned as annuals. Contrary to theoretical expectations, the species had low rates of juvenile mortality, and did not exhibit the demographic specializations of weedy or colonizing species. It is suggested that the annual habit in this case may be adaptive in sites with continual disturbance, such as that due to pocket gopher activity.     

Density-dependent regulation in populations of potato-colonizing aphids

Scarcity of long-term (over 30 years) data series represents a major challenge for an accurate estimation of the role of density-dependent processes in population regulation. We analyzed population densities of the wingless parthenogenic morphs of buckthorn aphid (BA), Aphis nasturtii Kaltenbach, potato aphid (PA), Macrosiphum euphorbiae (Thomas), and green peach aphid (GPA), Myzus persicae (Sulzer) from 1949 to 2003 for signs of density-dependent regulation. We found strong evidence of density-dependent regulation, with detection of density dependence being fairly consistent among the different statistical techniques. Direct density dependence was detected for the populations of all three species. There was also evidence of delayed density dependence for PA. The periodicity of population fluctuations for BA and GPA was 6.1 years and 3.9 years, respectively. The periodicity for PA was not explicit, being highly variable throughout the time series. Effects of density-independent weather factors were relatively minor compared to density-dependent regulation. The BA populations experienced a significant reduction in both density and annual oscillations starting in 1995, while GPA populations experienced a similar reduction in 1991. No such change was apparent for PA. The most likely explanation for the observed phenomenon is a change in the composition of the lady beetle community following the establishment of two alien coccinellid species, and/or changes in insecticide use by commercial growers in the area of the study.     

Hypothetical role of men's sexual activity in the regulation of human sex ratio: Analysis of the sex ratio of post-war abandoned children

The regulation of the sex ratio at birth in human species remains poorly understood. After wars, a shift of the sex ratio in favor of men is always observed. Among the different hypothesis to explain this observation, one is to consider that Y-bearing spermatozoids have a weight advantage following insemination and that X-bearing spermatozoids, heavier, are more time-resistant. Following these observations, frequent sex may favor the birth of boys, whether infrequent sex may favor the birth of girls.Sustaining this sperm weight hypothesis, I report here that in France, after the two world wars, there has been an increase of abandoned illegitimate children with a significant shift of the sex ratio in favor of men. These observations may reflect an increase in illegitimate birth and indirectly an increase of men paternity.     

Density-dependent self-fertilization and male versus hermaphrodite siring success in an androdioecious plant

Models of mating-system evolution emphasize the importance of frequency-dependent interactions among mating partners. It is also known that outcross siring success and the selfing rate in self-compatible hermaphrodites can be density dependent. Here, we use array experiments to show that the mating system (i.e., the outcrossing rate) and the siring success of morphs with divergent sex allocation strategies are both density dependent and frequency dependent in androdioecious populations of the wind-pollinated, annual plant Mercurialis annua. In particular, the outcrossing rate is a decreasing function of the mean interplant distance, regulated by a negative exponential pollen fall-off curve. Our results indicate that pollen dispersed from a male inflorescence are over 60% more likely to sire outcrossed progeny than equivalent pollen dispersed from hermaphrodites, likely due to the fact that males, but not hermaphrodites, disperse their pollen from erect inflorescence stalks. Because of this difference, and because males of M. annua produce much more pollen than hermaphrodites, the presence of males in the experimental arrays reduced both the selfing rate and the outcross siring success of hermaphrodites. We use our results to infer a density threshold below which males are unable to persist with hermaphrodites but above which they can invade hermaphroditic populations. We discuss our findings in the context of a metapopulation model, in which males can only persist in well-established populations but are excluded from small, sparse populations, for example, in the early stages of colonization.     

Variable variation: annual and seasonal changes in offspring sex ratio in a bat

Many organisms produce offspring with sex-ratios that deviate from equal numbers of males and females, and numerous adaptive explanations have been proposed. In some species, offspring sex-ratio varies across the reproductive season, again with several explanations as to why this might be adaptive. However, patterns for birds and mammals are inconsistent, and multiple factors are likely involved. Long-term studies on a variety of species may help untangle the complexity. I analyzed a long-term data set on the variation in offspring sex-ratio of the big brown bat, Eptesicus fuscus, a temperate-zone, insectivorous species. Sex ratio varied seasonally, but only in some years. Births early in the season were significantly female biased in years in which parturition occurred relatively early, but not in years with late parturition. Survival of female pups increased with earlier median birth date for the colony, and early-born females were more likely to survive and reproduce as one-year olds, compared to later-born pups. I argue that, due to the unusual timing of reproductive activities in male and female bats that hibernate, producing female offspring early in the year increases their probability of reproducing as one year olds, but this is not the case for male offspring. Thus, mothers that can give birth early in the year, benefit most by producing a female pup. The relative benefit of producing female or male offspring varies depending on the length of the growing season and thus the time available for female pups to reach sexual maturity. This suggests that not only does sex-ratio vary seasonally and among years, depending on the condition of the mother and the environment, but also likely varies geographically due to differences in season length.     

Offspring sex ratio under inbreeding and outbreeding in a gynodioecious plant

Silene vulgaris is a gynodioecious plant native to Eurasia and now found throughout much of North America. Using hermaphrodite plants from three geographic regions (Stamford, NY; Broadway,VA; and Giles Co., VA) and four local populations within each region, we employed a hierarchical crossing design to explore the geographic structure of sex determining genes. Sex determination in this species is cytonuclear involving multiple cytoplasmic male sterility and nuclear restorer loci. Due to dominance effects within nuclear restorer loci, self-fertilization of hermaphrodites heterozygous at restorer loci should produce some homozygous recessive female offspring. Female offspring may also result from outcrossing among related individuals. At greater geographic and genetic distances, mismatches between cytoplasmic and nuclear sex determining genes should also produce high frequencies of female offspring if coevolution between cytoplasmic and nuclear sex determining alleles occurs independently among widely separated populations. We found evidence of dominance effects among nuclear restorer loci but no evidence of nuclear-cytoplasmic mismatches at the regional level. Of 63 maternal lines, 55 produced at least one female offspring when self-fertilized. Outcrossing within populations produced significantly fewer female offspring than self-fertilization. Outcrossing among regions produced the lowest proportion of female offspring, significantly fewer than outcrossing among populations within regions. Regions responded differently to among-region outcrossing with pollen donors from the two Virginia regions producing far fewer female offspring with New York dams than crosses among New York populations. These results indicate that nuclear restoration is complex, involving multiple loci with epistatic interactions and that most hermaphrodites in nature are heterozygous at one or more restorer locus. Further, regional differences in restorer frequencies indicate significant genetic structure for sex determining genes at large geographic scales, perhaps reflecting invasion history.     

Density-dependent dispersal promotes female-biased sex allocation in viscous populations

A surprising result emerging from the theory of sex allocation is that the optimal sex ratio is predicted to be completely independent of the rate of dispersal. This striking invariance result has stimulated a huge amount of theoretical and empirical attention in the social evolution literature. However, this sex-allocation invariant has been derived under the assumption that an individual''s dispersal behaviour is not modulated by population density. Here, we investigate how density-dependent dispersal shapes patterns of sex allocation in a viscous-population setting. Specifically, we find that if individuals are able to adjust their dispersal behaviour according to local population density, then they are favoured to do so, and this drives the evolution of female-biased sex allocation. This result obtains because, whereas under density-independent dispersal, population viscosity is associated not only with higher relatedness—which promotes female bias—but also with higher kin competition—which inhibits female bias—under density-dependent dispersal, the kin-competition consequences of a female-biased sex ratio are entirely abolished. We derive analytical results for the full range of group sizes and costs of dispersal, under haploid, diploid and haplodiploid modes of inheritance. These results show that population viscosity promotes female-biased sex ratios in the context of density-dependent dispersal.     

Density-dependent regulation of breeding success in the Red-crested Pochard Netta rufina

Capsule: Numbers of breeding Red-crested Pochard Netta rufina in the Czech Republic were positively correlated with reproductive success in the previous year, yet annual productivity, declined with increasing numbers of females at the beginning of the breeding season, suggesting local limiting factors on this population.

Aims: To test the hypothesis that long-term increases in numbers of a breeding population of Red-crested Pochard Netta rufina can be explained by the productivity (numbers of ducklings fledged) in the previous year. We also test for the potential adverse effect of population size on population productivity in the current year.

Methods: Data from 2004 to 2016 from South Bohemia, Czech Republic, were analysed using TRends and Indices for Monitoring Data software to investigate changes in numbers of adults, numbers of females at the start of the breeding season and number of broods produced.

Results: There was a significant increase in the numbers of adults and females at the beginning of each breeding season. Numbers of adults in May positively correlated with productivity (numbers of ducklings fledged) in the previous breeding season, but relative productivity (broods per adult female) in the current year was negatively correlated with females abundance at the beginning of the breeding season.

Conclusions: The study identifies density-dependent regulation as a factor in the population productivity of Red-crested Pochards using intensively managed fishponds as a breeding ground.     

A model for the dynamics of an annual plant population

A model for the dynamics of a single species population of plants is proposed and its use demonstrated by the analysis of a simple example. The model incorporates the effects of microsite variation by allowing for individual differences in growth and death rates within each season. We demonstrate that an increase in the variance in individual growth rates may increase both the chances that a plant population will persist and the equilibrium size of that population. We also show that even if size-dependent death is occurring, it may not have a significant effect on the shape of the size frequency distribution. An extension of the model to multispecies communities of plants suggests an experimental procedure to determine whether competition is responsible for excluding a particular plant species from a community that appears otherwise to be suitable. A more detailed analysis of the model for a two-species community produces conditions for competitive coexistence reminiscent of those from the Lotka-Volterra competition equations. Another extension suggests that selection will favor those genotypes that maximize the product of germination probability and mass of seeds produced, if survivorship and growth are not substantially altered. Finally, an analog to r- and K-selection theory for animal populations is developed. Selection in low-density populations favors increasing growth rate, and in high-density populations favors minimizing the effect of neighbors on one's own growth rate.     

荒漠一年生植物小画眉草的种群动态调节与模拟

以分布在沙波头沙漠试验研究站人工固沙植被中的一年生植物小画眉草为例,研究了荒漠一年生植物不同时间尺度上的种群动态与调节机制,试验结果表明：（1）一年生植物小画眉草种群的续存与其种子库中种子的分批萌发对策密切相关;（2）同一个生长季内,制约小画眉草种群数量变化的关键因子随降水条件的不同而变化,即当环境适宜度较小时（干旱胁迫）,非生物因子（降水）限制种群数量;当环境适宜度较大时,密度依赖的竞争作用调节种群大小;（3）在较长的生态时间尺度上土壤特性（结皮厚度,养分含量）也是影响小画眉草种群动态的一个重要因素;（4）小画眉草种群的存活曲线属于C型,说明该种是较为典型的r－对策者。     

Density-dependent regulation of natural and laboratory rotifer populations

Density-dependent regulation of abundance is fundamentally important in the dynamics of most animal populations. Density effects, however, have rarely been quantified in natural populations, so population models typically have a large uncertainty in their predictions. We used models generated from time series analysis to explore the form and strength of density-dependence in several natural rotifer populations. Population growth rate (r) decreased linearly or non-linearly with increased population density, depending on the rotifer species. Density effects in natural populations reduced r to 0 at densities of 1–10 l^–1 for 8 of the 9 rotifer species investigated. The sensitivities of these species to density effects appeared normally distributed, with a mean r=0 density of 2.3 l^–1 and a standard deviation of 1.9. Brachionus rotundiformis was the outlier with 10–100× higher density tolerance. Density effects in laboratory rotifer populations reduced r to 0 at population densities of 10–100 ml^–1, which is 10⁴ higher than densities in natural populations. Density effects in laboratory populations are due to food limitation, autotoxicity or to their combined effects. Experiments with B. rotundiformis demonstrated the absence of autotoxicity at densities as high as 865 ml^–1, a much higher density than observed in natural populations. It is, therefore, likely that food limitation rather than autotoxicity plays a major role in regulating natural rotifer populations.     

Models of an annual plant population with a seedbank

Two models are set up, in terms of difference equations, for a population of a species of annual plant, of which a proportion of the seeds remain dormant for at least one year. Analytical and numerical results are presented demonstrating the effect of the seedbank on the return to equilibrium of the plant population following an initial disturbance. The seedbank can exert both a damping (stabilising) effect, speeding up oscillatory return, and an inertial (destabilising) effect, slowing down exponential return to the equilibrium population.