Different female mating rates in different populations do not reflect the benefits the females gain from polyandry in the adzuki bean beetle

The question why females in many species mate with several males (polyandry) has engaged the interest of evolutionary biologists for many years, and many studies have been conducted on the nature of the benefits that the females gain from polyandry. To understand the variation of female mating rates among species and populations it is indispensable to test the prediction that females of more polyandrous populations experience larger fitness benefit than those of less polyandrous populations. We compared the fitness components of two strains of the adzuki bean beetle Callosobruchus chinensis that have genetically different female mating rates. We measured the number of hatched eggs of once-copulated females and twice-copulated females in each strain. The statistical interaction for the number of hatched eggs between the number of matings and strains was determined. The increase in the number of hatched eggs is larger for the lower mating-rate strain than for the higher mating rate strain. This means that females of the lower mating-rate strain would have larger fitness gain from polyandry than those of the higher mating-rate strain. The actual mating rates of females did not reflect female interests in adzuki bean beetles, suggesting they are affected by sexual conflict.     

Modeling the interaction of physiological time, seasonal weather patterns, and delayed mating on population dynamics of codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae)

The effect of delaying female mating on population growth in codling moth (Cydia pomonella (L.)) was found to act on a physiological time (degree-day) basis and was predictable using a simple quadratic equation. When combined with previous work on degree-day based mortality, we were able to evaluate how the magnitude of population reduction and survival varied between sites, years, and generations at locations in California, Michigan, Pennsylvania and Washington states. In general, reductions in population growth associated with females mating 1–3 days after emergence were greater in warmer areas and during warmer times of the year. In any given year and location, the temperature profiles during peak flight were crucial in determining the population reductions, but over an 11-year period, the average seasonal temperature profile was more important. During the overwintering generation, conditions were relatively mild in all locations and only minor differences were observed in population growth rates between locations. Populations experiencing 1–3 days delay in female mating were reduced 8, 19 and 32 % compared to populations experiencing no delay, respectively. During the first summer generation, population reductions doubled compared to those seen in the overwintering generation. During the second summer flight, reductions in population growth rate at the three cooler locations decreased, while they increased in the warmer California location. Overall, the results show delayed mating can help understand how population growth is related to environmental conditions experienced naturally by insect populations and will help guide studies of the mechanisms of mating disruption, a technique used for pest suppression in agricultural and forest systems.     

Male-female interactions in Drosophila melanogaster: model with one-locus and three-alleles

We develop a fertility model of fitness that is general in that it does not assume that the fitnesses of the mating combinations are symmetrical or that they are additive or multilicative (i. e., that they can be inferred from fitnesses of the two genotypes involved in a mating). %he model considers one locus with three alleles. An experimental test with Drosophila rnelanogaster confirms that the fitnesses of the mating types depart from both additivity (or multiplicativity) and symmetry although this last property is of no consequence for the development of analytical models). urnerical simulations yield the same, or very nearly the same, equilibrium freuencies as the analytical model, independently of whether or not Hardy-Weinberg equilibrium Trequencies are assumed at the beginning of each selection cycle.     

Population and species variation of minisatellite DNA in Plantago

Three Plantago species were surveyed for within- and between-population variation at DNA sequences detected with the M13 minisatellite probe. The levels and patterns of variation detected by this probe correspond to those expected from the mating systems of the species. The highly-selfing species P. major has a relatively low variability of minisatellite sequences within populations and considerable differentiation between populations. The outcrossing species P. lanceolata exhibits higher minisatellite variability within populations and moderate differentiation between populations. P. coronopus, with a mixed mating system, has levels of variation intermediate between P. major and P. lanceolata. The levels of variation within and between populations corresponds, in general, to the levels of allozyme variation determined in an earlier study. Mating system and population structure appear to have a major influence on M13-detected fragment variation.     

A Two-Stage Test for Distinguishing Random,Pseudo-Random and Nonrandom Mating Populations

There is no such an implication that a population in Hardy-Weinberg equilibrium must have undergone random mating. Therefore, it is unequivocal that the usual tests for “Hardy-Weinberg equilibrium” are indeed tests for “random union of gametes” but not for “random mating”. In this paper, utilizing population characteristics expressed in equilibrium state (equilibrium or disequilibrium) and mating behavior (random or nonrandom), a two-stage testing procedure for distinguishing random, pseudo-random and nonrandom mating populations is proposed. At the first stage, a population is tested for Hardy-Weinberg equilibrium. If insignificant result (i.e., in equilibrium) is obtained, then to a second stage the population is further tested for mating behavior. Random mating-pairs data are needed here for analysis instead of random individuals for usual Hardy-Weinberg equilibrium tests. Since distinguishing the three types of mating populations depends on the combined results of two stages, the probability of correct determination of the two-stage tests is discussed by simulation studies.     

植物种群交配系统、亲本分析以及基因流动研究

 在过去的二十年中,有关植物种群交配与散布过程的研究与日俱增,重点集中在利用母本子代系列(Maternal progeny arrays)来估计种群间自交与异交的相对比例,种群的花粉散布与雄性育性变异的模式。早期的研究主要依靠排除法来确定亲本,但几乎同时也意识到基因流动事件几乎是检测不到的。在大多数估计中,难以做到为大多数非迁移子代确定唯一的亲本。因此,基因流动与雄性育性的最大似然性方法得以引入该领域的研究。本文介绍了用单位点和多位点模型来估计家系与种群自交与异交的相对频率,着重阐述了目前可用于亲本分析与基因流的估计方法。最后介绍了我们对木根麦冬交配系统与亲本分析的研究以及亲本分析将来的研究方向。     

Evolution of pre- and post-copulatory traits in female Drosophila melanogaster as a correlated response to selection for resistance to cold stress

Exposure to low temperatures reduces gamete viability and fecundity in females of insect species like Drosophila. Hence, adaptation to cold stress can in principle involve modifications in reproductive traits in females. Studies on resistance to cold stress have mostly addressed the evolution of adult survivorship post cold shock. Very few studies have addressed the evolution of reproductive traits in females in response to cold stress. We have successfully selected replicate populations of Drosophila melanogaster for resistance to cold shock. After 50 generations of selection, we investigated pre- and post-copulatory traits i.e. mating latency, copulation duration, mating frequency and progeny production in female flies exposed to cold shock or control conditions. Post cold shock, females from the selected populations were better at recovery in terms of mating latency, mating success, and progeny production relative to females from the control populations. Performance of the two types of females was not different under control conditions. These findings clearly indicate that adaptation to cold stress involves rapid modification of the reproductive traits.     

Genetic effects of different mating modes in Sinocalycanthus chinensis (Cheng et S.Y. Chang) Cheng et S.Y. Chang,an endangered species endemic to Zhejiang Province,China

The tertiary relict plant Sinocalycanthus chinensis, endemic to Zhejiang province with small populations and fragmented distribution, is an endangered plant in China. A relatively high species-level genetic diversity and low population-level genetic diversity exist in this species, and large genetic differentiation exists between two main populations with significant geographical isolation. Based on a previous artificial simulation mating test, amplified fragment length polymorphism (AFLP) marker was used in the present study to assess genetic diversity of filial generation colonies generated by different mating modes and to clarify genetic effects of various mating modes. The filial generation colonies generated by natural pollination and by geitonogamy were found to be similar to each other in terms of their relatively low genetic parameters and minimum genetic differentiation. This indicated that under natural conditions, selfing might occur at a higher proportion leading to the low genetic diversity within the population. The degree of genetic diversity of the filial generation colonies generated by outbreeding was highest, followed by inbreeding (xenogamy) and selfing (geitonogamy). Moreover, genetic differentiation between filial generation colonies generated by selfing (geitonogamy) and inbreeding (i.e., xenogamy) was much smaller than that between filial generation colonies generated by selfing and outbreeding. The results indicated that the genetic effect of outbreeding was much more dominant than other mating modes. Therefore, artificially promoting outbreeding between the two isolated geographically populations contributed to the enhancement of genetic diversity in populations in S. chinensis.     

Male display rate reveals ejaculate characteristics in the Trinidadian guppy,Poecilia reticulata.

Previous work (Reynolds and Gross 1992) has demonstrated that mate choice enables female guppies, Poecilia reticulata, to obtain heritable benefits (''good genes'') for their offspring. It is not yet known whether males also signal their functional fertility, that is, their viability as fathers, to female guppies. Our study tested the hypothesis that a male''s behaviour reveals characteristics of his ejaculate. We uncovered a strong correlation between display rate and sperm number in male guppies derived from two wild Trinidadian populations. This result indicates that information on the functional fertility of potential mates is available to females. However, our investigation also found a significant relationship between the frequency of sneaky mating and sperm number. Female guppies in wild populations, particularly those experiencing a high predation regime, are constantly subjected to sneaky mating attempts. As the males responsible for these sneaky matings have highest reserves of sperm they may have the capacity to undermine female choice. The implications of these findings for sperm competition are discussed.     

Separable models for age-structured population genetics

This paper is concerned with the applications of nonlinear age-dependent dynamics to population genetics. Age-structured models are formulated for a single autosomal locus with an arbitrary number of alleles. The following cases are considered: a) haploid populations with selection and mutation; b) monoecious diploid populations with or without mutation reproducing by self-fertilization or by two types of random mating. The diploid models do not deal with selection. For these cases the genic and genotypic frequencies evolve towards time-persistent forms, whether the total population size tends towards exponential growth or not.     

Sex-biased movement in the guppy (<Emphasis Type="Italic">Poecilia reticulata</Emphasis>)

The movement strategies of birds and mammals are often closely linked to their mating system, but few studies have examined the relationship between mating systems and movement in fishes. We examined the movement patterns of the guppy (Poecilia reticulata) in the Arima river of Trinidad and predicted that sexual asymmetry in reproductive investment would result in male-biased movement. Since male guppies maximize their reproductive success by mating with as many different females as possible, there should be strong selection for males to move in search of mates. In agreement with our prediction, the percentage of fish that emigrated from release pools was higher for males than females (27.3% vs. 6.9%, respectively). Sex ratio was highly variable among pools and may influence a male's decision to emigrate or continue moving. We also detected a positive relationship between body length and the probability of emigration for males and a significant bias for upstream movement by males. Among the few females that did emigrate, a positive correlation was observed between body length and distance moved. Sex-biased movement appears to be related to mating systems in fishes, but the evidence is very limited. Given the implications for ecology, evolution, and conservation, future studies should explicitly address the influence of sex and mating systems on movement patterns.     

Barriers to outcrossing success in the primarily self-fertilizing clam shrimp, Eulimnadia texana (Crustacea, Branchiopoda)

Abstract. The expected proportion of males in androdioecious populations (those comprised of males and hermaphrodites) largely depends on the fertilization opportunities of males. If male mating opportunities are low due to restricted access to hermaphroditic eggs, then populations will be hermaphrodite-biased. Hermaphrodites have two mechanisms available to limit male mating success: (1) pre-mating barriers to outcrossing, in which hermaphrodites choose not to pair with males and (2) post-mating barriers to outcrossing, in which hermaphrodite sperm has greater access to eggs than male sperm. In this study, we measured male mating success in the androdioecious clam shrimp Eulimnadia texana when pre-mating barriers to outcrossing were removed. These branchiopod crustaceans are small (5–8 mm), filter feeders that live in ephemeral pools in the deserts of the southwestern United States. Using genetic markers, we measured male mating success in laboratory experiments in two populations of these shrimp. We correlated mating success with clasping time, clasping during egg transfer, and male thrusting during egg transfer. Males fertilized an average of 24–40% of the hermaphrodites' eggs. Outcrossing success was positively correlated with clasping duration, and was nearly an order of magnitude higher for males thrusting during egg transfer relative to thrusting at other times during pairing. Because these estimates of mating success were similar to previously reported estimates (in which both pre- and post-mating barriers to outcrossing were potentially important), we deduced that pre-mating barriers to outcrossing do not greatly decrease male outcrossing success in E. texana ; the low fertilization (25–50% of available eggs) by males is thus due to post-mating barrier(s) to outcrossing.     

Mating pattern variability among western toad (Bufo boreas) populations

Summary Breeding ecology and mating patterns of the western toad, Bufo boreas, were examined in three large explosively breeding populations in the Oregon Cascade Mountains. Two mating patterns occurred variably within and among the three populations. First, a large male mating advantage was observed in two of the three populations when data from all the days of data collection were combined. When each day of breeding was analyzed separately, there was a large male mating advantage on 3 of 5 days at one population, and mating was random on all days at the two other populations. The second mating pattern, positive assortative mating by size, was observed at two of the three populations. This pattern was found on separate days of breeding as well as when data from all days were combined at one population, and on only one day of breeding at the second population.In a survey of anuran amphibian mating patterns, intraspecific variation was found in 13 of 15 species, including the present study of the western toad. Intrapopulation variation in mating patterns among breeding years has been observed in 5 of 8 anurans, whereas within-site, within-year mating pattern variation has only been reported for the western toad. These results strongly suggest that anuran mating patterns are frequently neither species-specific nor population-specific attributes. Variable mating patterns were most commonly observed in explosively-breeding anurans. Explosive breeders may be susceptible to variable mating patterns because they may be more sensitive to fluctuations in environmental conditions, demographic parameters, and the intensities of intrasexual competition and mate choice.     

红树植物木榄(Bruguiera gymnorrhiza)种群的交配系统

利用等位酶多态位点作遗传标记,采用水平切片淀粉凝胶电泳技术检测了我国福建浮宫（24°24′）、广东深圳福田（22°32′）、广西山口合浦（21°28′）和海南东寨港（19°51′）4个红树植物分布区内木榄种群的基因型,进而利用多位点异交率估测程序（MLT）估测其交配系统。所用多态酶位点在浮宫、广西、海南均为Mdh-1、Mdh-2、Me-1,在深圳为Mdh-1、Mdh-2、Aat-1、Aat-2。结果表明各种群多态位点异交率之间有很大差别,浮宫种群最大,为0.845;海南种群最小,为0.267。多位点异交率与单位点异交率平均值的差值显示出除广西种群为随机交配外,其余3个种群都存在轻度的双亲近交。因此,木榄属于混合交配类型,以杂交为主。这主要受植物种群密度、结构,传粉者适应性和活动能力以及是否自交亲和等因素的影响。     

INCIPIENT REPRODUCTIVE ISOLATION BETWEEN TWO SYMPATRIC MORPHS OF THE INTERTIDAL SNAIL LITTORINA SAXATILIS

The study of speciation in recent populations is essentially a study of the evolution of reproductive isolation mechanisms between sub-groups of a species. Prezygotic isolation can be of central importance to models of speciation, either being a consequence of reinforcement of assortative mating in hybrid zones, or a pleiotropic effect of morphological or behavioral adaptation to different environments. To suggest speciation by reinforcement between incipient species one must at least know that gene flow occurs, or have recently occurred, and that assortative mating has been established in the hybrid zone. In Galician populations of the marine snail Littorina saxatilis, two main morphs appear on the same shores, one on the upper-shore barnacle belt and the other in the lower-shore mussel belt. The two morphs overlap in distribution in the midshore where hybrids are found together with pure forms. Allozyme variation indicates that the two parental morphs share a common gene pool, although within shores, gene flow between morphs is less than gene flow within morphs. In this study, we observed mating behavior in the field, and we found that mating was not random in midshore sites, with a deficiency of heterotypic pairs. Habitat selection, assortative mating, and possibly sexual selection among females contributed to the partial reproductive isolation between the pure morphs. Sizes of mates were often positively correlated, in particular, in the upper shore, indicating size-assortative mating too. However, this seemed to be a consequence of nonrandom microdistributions of snails of different sizes. Because we also argue that the hybrid zone is of primary rather than secondary origin, this seems to be an example of sympatric reproductive isolation, either established by means of reinforcement or as a by-product to divergent selection acting on other characters.     

Pheromone trapping models for pest control: Effects of mating patterns and immigration

Several models are presented which examine pest population behaviour with the release of female sex pheromones for the attraction and annihilation of males. These models include male polygamy and female monogamy, various mating frequencies, delayed mating of females, immigration of one or all individual types, and differential survivorship of males and females. In all the models there are two steady states, a stable s.s. at the origin and an unstable s.s. in the positive domain for a given value of pheromone release rate. In all the models, control relies on the reduced ability of males to fertilize virgin females following trapping and male annihilation. As such, control is very sensitive to mating frequency, being very difficult when males mate frequently. Control is also very difficult with the immigration of even a moderate number of fertilized females. Control is much easier when mating is delayed, especially if survivorship is low, or with density dependent population regulation.     

Sex, sex-ratios, and the dynamics of pelagic copepod populations

I examine how the population biology of pelagic copepods depends on their mating biology using field data and a simple demographic model. Among calanoid copepods, two distinct patterns emerge. Firstly, copepods that lack seminal receptacle and require repeated mating to stay fertilized have near equal adult sex ratios in field populations. Winter population densities are orders of magnitude less than the critical population density required for population persistence, but populations survive winter seasons as resting eggs in the sediment. Population growth in these species is potentially high because they have on average a factor of 2 higher egg production rates than other pelagic copepods. Secondly, other copepods require only one mating to stay fertile, and populations of these species have strongly female-skewed adult sex-ratios in field populations. Resting eggs have not been described within this group. Winter population sizes are well predicted by the critical density required for population persistence which, in turn, is closely related to the body-size-dependent mate-search capacity. Thus, the different requirements for mating lead in the first case to a more opportunistic reproductive strategy that implies rapid colonization of the pelagic during productive seasons, and in the second case to a strategy that allows maintenance of a pelagic populations during unproductive seasons. Positive density dependent population growth during periods of low population density (‘Allee effect’) amplifies population density variation during winter into the subsequent summer, thus explaining why summer densities appear to depend more on winter densities than on current growth opportunities in pelagic copepods.     

Non-Mendelian segregation influences the infection biology and genetic structure of the African tree pathogen Ceratocystis albifundus

The African fungal tree pathogen, Ceratocystis albifundus, undergoes uni-directional mating type switching, giving rise to either self-fertile or self-sterile progeny. Self-sterile isolates lack the MAT1-2-1 gene and have reduced fitness such as slower growth and reduced pathogenicity, relative to self-fertile isolates. While it has been hypothesized that there is a 1:1 ratio of self-fertile to self-sterile ascospore progeny in relatives of C. albifundus, some studies have reported a significant bias in this ratio. This could be due to the fact that either fewer self-sterile ascospores are produced or that self-sterile ascospores have low viability. We quantified the percentage of self-sterile and self-fertile ascospores from ascospore masses in C. albifundus using real-time PCR. Primers were designed to distinguish between spores that contained the MAT1-2-1 gene and those where this gene had been deleted. A significant bias towards the self-fertile mating type was observed in all single ascospore masses taken from sexual structures produced in haploid-selfed cultures. The same result was observed from a disease outbreak situation in an intensively managed field of cultivated native trees, and this was coupled with very low population diversity in the pathogen. This was in contrast to the results obtained from ascospore masses taken from the crosses performed under laboratory conditions or ascomata on native trees in a non-disease situation, where either self-fertile or self-sterile ascospores were dominant. The results suggest that reproductive strategies play a significant role in the infection biology and genetic structure of C. albifundus populations.     

Breeding systems of hermaphroditic and gynodioecious populations of the colonizing species Trifolium hirtum All. in California

Summary A multilocus procedure was used to estimate outcrossing rates in ten roadside populations of Trifolium hirtum in California. Three groups of populations were studied: cultivars, hermaphroditic, and gynodioecious (sexually dimorphic) populations. The multilocus outcrossing rate (t_m) varied from 0.05 to 0.43 among populations. Population level t_m estimates were significantly correlated with the observed heterozygosity in gynodioecious populations but not in hermaphroditic populations. The outcrossing rate of hermaphrodites and females was estimated in three gynodioecious populations; the estimates of t_m varied from 0.09 to 0.23 for hermaphrodites and from 0.73 to 0.80 for females. The distribution of outcrossing rates in gynodioecious populations is bimodal. Our results indicate that for the levels of selfing observed among hermaphrodites, inbreeding depression is likely to be a major factor in the maintenance of females in gynodioecious populations.     

Mating disruption or mass trapping? Numerical simulation analysis of a control strategy for lepidopteran pests

I have constructed a simulation model applicable to both mass trapping and mating disruption for lepidopteran insect pests. The basic structure of the model is based on mass trapping model proposed by Knipling and McGuire (Agric Info Bull 308:1–20 1966), but this was modified to include mechanistic competition among females and lures. Several new implications are derived from the model. (1) Long-living pests are hard to control. (2) Protandry does not improve control efficiency for pests with low survival rates. (3) Sexual communication across large distances is more difficult to control than that across a short range. (4) There is an upper limit to improvement which can be achieved by increasing the number of pheromone traps. (5) Improving the catching efficiency of traps does not improve mating suppression although improvement of lure efficiency does substantially. The last implication, in particular, has practical importance. If an efficient lure attracts males or inhibits their ability to locate females and mate, mating disruption works as well as mass trapping with the same number of lures. In such circumstances mating disruption should be preferred to mass trapping because the former does not incur the cost of the trapping devices. Mass trapping should, however, be considered in preference to mating disruption if the lure is not efficient enough and no other methods are available. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.