Predominant Mode Of Reproduction Research Articles

Geographical Parthenogenesis in Alpine and Arctic Plants

The term "Geographical parthenogenesis" describes the phenomenon that asexual organisms usually occupy larger and more northern distribution areas than their sexual relatives, and tend to colonize previously glaciated areas. Several case studies on alpine and arctic plants confirm the geographical pattern, but the causal factors behind the phenomenon are still unclear. Research of the last decade in several plant families has shed light on the question and evaluated some of the classical evolutionary theories. Results confirmed, in general, that the advantages of uniparental reproduction enable apomictic plants to re-colonize faster in larger and more northern distribution areas. Associated factors like polyploidy seem to contribute mainly to the spatial separation of sexual and asexual cytotypes. Ecological studies suggest a better tolerance of apomicts to colder climates and temperate extremes, whereby epigenetic flexibility and phenotypic plasticity play an important role in occupying ecological niches under harsh conditions. Genotypic diversity appears to be of lesser importance for the distributional success of asexual plants. Classical evolutionary theories like a reduced pressure of biotic interactions in colder climates and hence an advantage to asexuals (Red Queen hypothesis) did not gain support from studies on plants. However, it is also still enigmatic why sexual outcrossing remains the predominant mode of reproduction also in alpine floras. Constraints for the origin of apomixis might play a role. Interestingly, some studies suggest an association of sexuality with abiotic stresses. Light stress in high elevations might explain why most alpine plants retain sexual reproduction despite other environmental factors that would favor apomixis. Directions for future research will be given.

Substantial genetic mixing among sexual and androgenetic lineages within the clam genus Corbicula

“Occasional” sexuality occurs when a species combines clonal reproduction and genetic mixing. This strategy is predicted to combine the advantages of both asexuality and sexuality, but its actual consequences on the genetic diversity and species longevity are poorly understood. Androgenesis, a reproductive mode in which the offspring inherits its entire nuclear genome from the father, is often reported as a strictly clonal reproductive mode. Androgenesis is the predominant reproductive mode within the hermaphroditic, invasive lineages of the mollusk genus Corbicula. Their ability to reproduce clonally through androgenesis has been determinant in their invasive success, having colonized during the 20th century American and European freshwater systems, where they became notorious invaders with a widespread, global distribution. However, in androgenetic Corbicula clams, occasional genetic mixing between distinct lineages has also been observed when the sperm of one lineage fertilizes the oocyte of another one. Because of these occasional introgressions, the genetic relationships between Corbicula species remained unclear, and the biogeographic origins of the invasive androgenetic lineages have been challenging to identify. To address these issues, we analyzed the patterns of allele sharing for several nuclear and mitochondrial molecular markers among Corbicula individuals collected across both the native and invasive range. Our results show the occurrence of an allelic pool encompassing all Corbicula freshwater species worldwide, including sexual and androgenetic ones, which highlights the substantial genetic mixing within this genus. However, the differences in allele sharing patterns between invasive lineages, and the low diversity within each lineage, suggest recent, distinct biogeographic origins of invasive Corbicula androgenetic lineages. Finally, the polyploidy, high heterozygosity, and hybrid phenotypes and genotypes found in our study probably originated from hybridization events following egg parasitism between distinct Corbicula lineages. This extensive cross-lineage mixing found in Corbicula may generate nuclear diversity in an otherwise asexually reproducing species.

Distinguishing Homokaryons and Heterokaryons in Medicinal Polypore Mushroom Wolfiporia cocos (Agaricomycetes) Based on Cultural and Genetic Characteristics.

The sclerotia of Wolfiporia cocos are a kind of traditional medicine and food with excellent benefits and are widely used in China, Japan, and other Asian countries. The mating system of fungi is not only of practical importance for breeding but also has profound effects on genetic variability and molecular evolution. However, the lack of clamp connections in W. cocos increases the difficulty of research on mating systems. In this study, homokaryons and heterokaryons were distinguished by comparing the characteristics of culture, fruiting tests, and molecular markers, which was further demonstrated by k-mer analysis based on Illumina sequencing. Uninucleate, binucleate, and nuclei-free condition basidiospores of W. cocos were observed, and binucleate basidiospores were the most predominant. Brown-type colonies, slow growth rates in both PDA medium and sawdust substrate, and neutral pH after the growth of mycelia and unfruiting were found to be the morphological and growth characteristics of homokaryotic strains. Primers SSR37 and 38 were screened to identify homokaryons. K-mer analysis based on Illumina sequencing exhibited different heterozygous ratios for homokaryons and heterokaryons. The results revealed that pseudo-homothallism was the predominant mode of reproduction in the Chinese population of W. cocos, and heterothallism also existed in all probability. This study will be helpful for the cross-breeding of this precious medicinal mushroom and for understanding its evolution and population structure.

Variation over time in morphological phenotypes and reproductive behavior in a natural wild potato population from Tucumán, Argentina

The common potato, Solanum tuberosum ssp. tuberosum (tbr, 2n = 4x = 48, 4EBN), has a large number of wild relatives (known as “wild potatoes”) with wide geographic and ecological distribution in the Americas. They form a polyploid series with 2n = 2x to 6x (x = 12), have scarce chromosome differentiation, and can reproduce both sexually (by seeds) and asexually (by tubers and stolons). In nature, wild potatoes can be isolated by external and/or internal hybridization barriers; when incomplete, internal barriers allow for gene flow and introgression between sympatric populations. To develop appropriate strategies to in situ maintain the available genetic diversity and to capture a large part of this diversity for ex situ conservation, it is of utmost importance to understand the reproductive behavior of wild potatoes in their natural environments, and to explore possible over time changes. Information in this regard is lacking. Thus, an integrated analysis of morphological phenotypes and breeding relationships was carried out in a natural population from Tucuman province, Argentina. This population was sampled in two consecutive years following a field design and, ex situ, controlled crosses were carried out between plants derived from the sampled propagules. High morphological variability and absence of relationships between morphological phenotypes and breeding behavior was observed, in addition to changes in the predominant mode of reproduction (either sexual or asexual) over time. These results highlight the necessity of resampling wild potato populations and of applying the gene reservoir concept in natural populations -irrespective of individual plant morphological phenotypes- for both conservation and use of their genetic diversity.

Ginger and the beetle: Evidence of primitive pollination system in a Himalayan endemic alpine ginger (Roscoea alpina, Zingiberaceae).

The Himalayan endemic alpine genus Roscoea, like other members of ginger family, exhibits the combination of floral traits that would fit pollination by long distant foragers such as bees, birds or flies. We studied the pollination biology of Roscoea alpina, observed potential floral visitors and determined their foraging behaviour, visitation frequency and pollination efficiency, to seek evidence in support of the pollination syndrome hypothesis. We also measured the floral spectra of R. alpina flowers to evaluate if signals fit with the currently known framework for observed floral visitors. We found that R. alpina have autonomous selfing and pollinator-mediated crossing, but lack apomixis. We observed that a beetle (Mylabris sp.), and a moth (Macroglossum nycteris) visit the flowers of R. alpina for pollen and nectar feeding respectively. Our field observations, the stigmatic pollen count and fruit set data indicated that the visit by the beetle was legitimate, while that of the moth was illegitimate. Emasculated flowers visited by beetles set as many fruits and seeds/fruit as auto-selfed and naturally pollinated flowers, while emasculated flowers excluded from beetle visits did not set fruit and seed; indicating that a single visit of a beetle to the flowers of R. alpina can facilitate pollination. We found that flower spectral signal of R. alpina does not fit typical spectra previously reported for beetle or bee-visited flowers. Our results suggest that, to ensure reproductive success in alpine habitat, R. alpina has evolved autonomous selfing as a predominant mode of reproduction, while beetle pollination would promote genetic diversity of this plant species. The visitation of beetles to the flowers of R. alpina, despite floral signal mismatch with the classically associated beetle vision, suggests that a different visual processing may operate in this plant-pollinator interaction at high altitudes.

Genetic Structure of Populations of the Wheat Sharp Eyespot Pathogen Rhizoctonia cerealis Anastomosis Group D Subgroup I in China.

Sharp eyespot on wheat is caused by Rhizoctonia cerealis anastomosis group D subgroup I (AG-DI) and is an economically important stem-base disease of wheat in temperate regions worldwide. However, the understanding about the field population structure of R. cerealis is limited. In this study, the genetic structure of four wheat-infecting populations in China was investigated using six microsatellite markers characterized from the transcriptome data of R. cerealis AG-DI. A total of 173 unique genotypes were identified among 235 fungal isolates. Departure from Hardy-Weinberg equilibrium, a significant degree of inbreeding, and a significant deficit in heterozygotes indicated a nonrandom mating pattern. Combining the low to intermediate degrees of gametic disequilibrium, although with high genotypic diversity and low to moderate clonal fractions, sexual reproduction probably existed, but the asexual reproduction should be the predominant reproductive mode. Structural analysis showed three gene pools among the four populations, which indicated the existence of three evolutionary origins of R. cerealis AG-DI. The long-distance movement of contaminated material, especially the infected seed, might have caused the moderate gene flow among these populations, which was consistent with the high differentiation among these populations. Overall, the genetic characteristics of the populations suggested a moderate evolutionary potential for R. cerealis AG-DI in China.

Recent Introduction and Recombination in Colletotrichum acutatum Populations Associated with Citrus Postbloom Fruit Drop Epidemics in São Paulo, Brazil

Citrus crops in São Paulo State, Brazil, have been severely affected by postbloom fruit drop disease (PFD), which is caused by Colletotrichum acutatum. This disease leads to the drop of up to 100% of young fruits. Previous studies have assumed that this pathogen exhibits a clonal reproductive mode, although no population genetic studies have been conducted so far. Thus, the genetic structure of six C. acutatum populations from sweet orange orchards showing PFD symptoms was determined using nine microsatellite markers, enabling inference on predominant mode of reproduction. C. acutatum populations exhibit a nearly panmictic genetic structure and a high degree of admixture, indicating either ongoing contemporary gene flow at a regional scale or a recent introduction from a common source, since this pathogen was introduced in Brazil only very recently. Sharing haplotypes among orchards separated by 400 km suggests the natural dispersal of fungal propagules, with the possible involvement of pollinators. A significant population expansion was detected, which was consistent with an increase in host density associated with crop expansion toward new areas across the state. Findings of moderate to high levels of haplotypic diversity and gametic equilibrium suggest that recombination might play an important role in these pathogen populations, possibly via parasexual reproduction or a cryptic sexual cycle. This study provides additional tools for epidemiological studies of C. acutatum to improve prevention and management strategies for this disease.

Epidemiology of asexuality induced by the endosymbioticWolbachiaacross phytophagous wasp species: host plant specialization matters

Among eukaryotes, sexual reproduction is by far the most predominant mode of reproduction. However, some systems maintaining sexuality appear particularly labile and raise intriguing questions on the evolutionary routes to asexuality. Thelytokous parthenogenesis is a form of spontaneous loss of sexuality leading to strong distortion of sex ratio towards females and resulting from mutation, hybridization or infection by bacterial endosymbionts. We investigated whether ecological specialization is a likely mechanism of spread of thelytoky within insect communities. Focusing on the highly specialized genus Megastigmus (Hymenoptera: Torymidae), we first performed a large literature survey to examine the distribution of thelytoky in these wasps across their respective obligate host plant families. Second, we tested for thelytoky caused by endosymbionts by screening in 15 arrhenotokous and 10 thelytokous species for Wolbachia, Cardinium, Arsenophonus and Rickettsia endosymbionts and by performing antibiotic treatments. Finally, we performed phylogenetic reconstructions using multilocus sequence typing (MLST) to examine the evolution of endosymbiont-mediated thelytoky in Megastigmus and its possible connections to host plant specialization. We demonstrate that thelytoky evolved from ancestral arrhenotoky through the horizontal transmission and the fixation of the parthenogenesis-inducing Wolbachia. We find that ecological specialization in Wolbachia's hosts was probably a critical driving force for Wolbachia infection and spread of thelytoky, but also a constraint. Our work further reinforces the hypothesis that community structure of insects is a major driver of the epidemiology of endosymbionts and that competitive interactions among closely related species may facilitate their horizontal transmission.

Assessing the effects of light on differentiation and virulence of the plant pathogen Botrytis cinerea: characterization of the White Collar Complex.

Organisms are exposed to a tough environment, where acute daily challenges, like light, can strongly affect several aspects of an individual's physiology, including pathogenesis. While several fungal models have been widely employed to understand the physiological and molecular events associated with light perception, various other agricultural-relevant fungi still remain, in terms of their responsiveness to light, in the dark. The fungus Botrytis cinerea is an aggressive pathogen able to cause disease on a wide range of plant species. Natural B. cinerea isolates exhibit a high degree of diversity in their predominant mode of reproduction. Thus, the majority of naturally occurring strains are known to reproduce asexually via conidia and sclerotia, and sexually via apothecia. Studies from the 1970′s reported on specific developmental responses to treatments with near-UV, blue, red and far-red light. To unravel the signaling machinery triggering development – and possibly also connected with virulence – we initiated the functional characterization of the transcription factor/photoreceptor BcWCL1 and its partner BcWCL2, that form the White Collar Complex (WCC) in B. cinerea. Using mutants either abolished in or exhibiting enhanced WCC signaling (overexpression of both bcwcl1 and bcwcl2), we demonstrate that the WCC is an integral part of the mentioned machinery by mediating transcriptional responses to white light and the inhibition of conidiation in response to this stimulus. Furthermore, the WCC is required for coping with excessive light, oxidative stress and also to achieve full virulence. Although several transcriptional responses are abolished in the absence of bcwcl1, the expression of some genes is still light induced and a distinct conidiation pattern in response to daily light oscillations is enhanced, revealing a complex underlying photobiology. Though overlaps with well-studied fungal systems exist, the light-associated machinery of B. cinerea appears more complex than those of Neurospora crassa and Aspergillus nidulans.

Multiple gene genealogical analyses of a nematophagous fungus Paecilomyces lilacinus from China

Paecilomyces lilacinus is a geographically widespread nematophagous fungus and a promising biological control agent against plant parasitic nematodes. However, relatively little is known about its patterns of genetic variation through its broad geographic and ecological contexts. In this study, we analyzed the genetic variation of 2 virulence-associated genes (PLS and PLC) and 4 housekeeping gene fragments (ITS, RPB1, RPB2, and β-tubulin) among 80 P. lilacinus specimens collected from 7 locations in China. Various degrees of polymorphism and haplotype diversity were observed among the six gene fragments. However, no genetic differentiation was observed among the geographic populations, consistent with extensive gene flow among these geographic populations of P. lilacinus in China. Our analysis also suggested that clonal reproduction was the predominant mode of reproduction in natural populations of P. lilacinus.

Fragment production and recruitment ecology of the red alga Laurencia poiteaui in Florida Bay, USA

In 2000 and 2001, Laurencia poiteaui (Rhodophyta, Ceramiales) was the dominant organism in the patchy seagrass beds off Long Key in Florida Bay. Earlier research established that asexual fragmentation is the predominant mode of reproduction in this benthic drift macroalga, yet no studies documenting post-fragment success have been published to date. To better understand fragmentation, and ultimately recruitment of L. poiteaui, we examined fragment: 1) accumulation rates, 2) variability, 3) dispersal distances, and 4) attachment rates. In July 2000 and July 2001, the majority (>50%) of the algal fragment pool near Long Key was comprised of L. poiteaui. The length of these fragments ranged from 0.7 to 15.7cm, but >75% of the collected fragments were <6cm. Short dispersal distances (<7cm within beds of the seagrass Thalassia testudinum and<30cm over sand after 7days) and fast attachment rates (<48h) suggest that most fragments (3–4fragments/day/m2) recruit locally. Attachment success of L. poiteaui fragments was high in both laboratory and field (95.5% and 88.2%, respectively). Sand and silt decreased the attachment rate of L. poiteaui, but the presence of a terminal apical tip had no influence. The combination of high fragment production, slow dispersal rates, and rapid attachment rates of L. poiteaui ultimately suggests that vegetative fragmentation plays an important role in the abundance of this macroalga in the calm, shallow areas of southeastern Florida Bay.

Sex at the margins: parthenogenesis vs. facultative and obligate sex in a Neotropical ant

Geographic parthenogenesis is a distribution pattern, in which parthenogenetic populations tend to live in marginal habitats, at higher latitudes and altitudes and island-like habitats compared with the sexual forms. The facultatively parthenogenetic ant Platythyrea punctata is thought to exhibit this general pattern throughout its wide range in Central America and the Caribbean Islands. Workers of P.punctata from the Caribbean produce diploid female offspring from unfertilized eggs by thelytokous parthenogenesis, and mated females and males are rare. In contrast, workers in one colony from Costa Rica were incapable of thelytoky; instead mated workers produced all female offspring. Because sample sizes were very low in former studies, we here use microsatellite markers and explicit tests of thelytoky to examine the population genetic structure of ancestral and derived populations of P.punctata throughout the Caribbean and Central America. Populations from the Caribbean islands were fully capable of parthenogenesis, and population genetic signatures indicate that this is the predominant mode of reproduction, although males are occasionally produced. In contrast, the northernmost population on the mainland (Texas) showed signatures of sexual reproduction, and individuals were incapable of reproduction by thelytoky. Contrary to expectations from a geographic parthenogenesis distribution pattern, most parts of the mainland populations were found to be facultatively thelytokous, with population genetic signatures of both sexual and parthenogenetic reproduction.

Contemporary cryptic sexuality in Trypanosoma cruzi

Clonal propagation is considered to be the predominant mode of reproduction among many parasitic protozoa. However, this assumption may overlook unorthodox, infrequent or cryptic sexuality. Trypanosoma cruzi, which causes Chagas disease, is known to undergo non-Mendelian genetic exchange in the laboratory. In the field, evidence of extant genetic exchange is limited. In this study, we undertook intensive sampling of T. cruzi Discrete Typing Unit I in endemic eastern Colombia. Using Fluorescence-activated cell sorting, we generated 269 biological clones from 67 strains. Each clone was genotyped across 24 microsatellite loci. Subsequently, 100 representative clones were typed using 10 mitochondrial sequence targets (3.76 Kbp total). Clonal diversity among humans, reservoir hosts and vectors suggested complex patterns of superinfection and/or coinfection in oral and vector-borne Chagas disease cases. Clonal diversity between mother and foetus in a congenital case demonstrates that domestic TcI genotypes are infective in utero. Importantly, gross incongruence between nuclear and mitochondrial markers is strong evidence for widespread genetic exchange throughout the data set. Furthermore, a confirmed mosaic maxicircle sequence suggests intermolecular recombination between individuals as a further mechanism of genetic reassortment. Finally, robust dating based on mitochondrial DNA indicates that the emergence of a widespread domestic TcI clade that we now name TcI(DOM) (formerly TcIa/VEN(Dom)) occurred 23 000 ± 12 000 years ago and was followed by population expansion, broadly corresponding with the earliest human migration into the Americas.

Exploring symbiont management in lichens

Lichens are unique among fungal symbioses in that their mycelial structures are compact and exposed to the light as thallus structures. The myriad intersections of unique fungal species with photosynthetic partner organisms (green algae in 90% of lichens) produce a wide variety of diverse shapes and colours of the fully synthesized lichen thallus when growing in nature. This characteristic complex morphology is, however, not achieved in the fungal axenic state. Even under ideal environmental conditions, the lichen life cycle faces considerable odds: first, meiotic spores are only produced on well-established thalli and often only after achieving considerable age in a stable environment, and second, even then in vivo resynthesis requires the presence of compatible algal strains where fungal spores germinate. Many lichen species have evolved a way around the resynthesis bottleneck by producing asexual propagules for joint propagation of symbionts. These different dispersal strategies ostensibly shape the population genetic structure of lichen symbioses, but the relative contributions of vertical (joint) and horizontal (independent) symbiont transmission have long eluded lichen evolutionary biologists. In this issue of Molecular Ecology, Dal Grande et al. (2012) close in on this question with the lung lichen, Lobaria pulmonaria, a flagship species in the conservation of old growth forests. By capitalizing on available microsatellite markers for both fungal and algal symbionts, they show that while vertical transmission is the predominant mode of reproduction, horizontal transmission is demonstrable and actively shapes population genetic structure. The resulting mixed propagation system is a highly successful balance of safe recruitment of symbiotic clones and endless possibilities for fungal recombination and symbiont shuffling.

Evolutionary dynamics of mating system shifts in Arabidopsis lyrata

Outcrossing is the prevalent mode of reproduction in plants and animals despite its substantial costs, while selfing and mixed mating occur at much lower frequency. Comparative research on plants has demonstrated the lability of self-incompatibility, but there is little information about the transition on a within-species level from self-incompatibility to predominant selfing. We studied variation in mating system among 18 populations of Arabidopsis lyrata within a phylogenetic context to shed light on the evolution of selfing. Realized and potential mating systems were assessed by genetic analysis with microsatellite markers and hand-self-pollinations on 30 plants from each population. The fraction of self-incompatible plants in a population was highly correlated with the outcrossing rate, showing that the spread of self-compatibility is accompanied by or soon followed by an increase in the rate of selfing. The four predominantly selfing populations (outcrossing rates <0.25) fell into more than one phylogenetic cluster, suggesting that the transition to selfing occurred more than once independently. Hence, A. lyrata offers an opportunity for the comparative analysis of outcrossing as a predominant mode of reproduction in plants and of the causes of the shift to selfing.

Vegetative reproduction facilitates early expansion of Phyla canescens in a semi-arid floodplain

Phyla canescens (Verbenaceae, hereafter lippia) is an invasive perennial forb from South America that is now widespread in Australia, threatening the biological values of internationally significant wetlands (Ramsar listed) and the productivity of the grazing industry (Crawford 2008; Earl 2003). Under certain conditions, lippia can displace native herbaceous vegetation and form mono-specific stands, but little is known about the process of colonization and early invasion. Lippia has a dual reproductive strategy, recruiting via seedlings and vegetative fragments in periods following inundation (Macdonald 2008). Determining the relative contribution of vegetative and sexual reproduction for lippia across environmental gradients is important for management action—if spread is primarily by vegetative fragments then control efforts should be focussed on managing vectors (e.g. movement of stock and vehicles), whereas if spread is mostly by seeds then management should focus on reducing the conditions favourable to seed production and seedling establishment. The predominant reproductive mode also has implications for the selection of biocontrol agents. A comparison of lippia in Australia with native range populations found that Australian populations had evolved to invest more energy in sexual reproduction than in the native range (Xu et al. 2009). Genetic analysis found that the importance of vegetative and sexual reproduction is highly variable among populations in Australia (Fatemi and Gross, unpublished data). In some cases, populations exhibited extremely low diversity, with single clones covering distances of up to 100 m and in others, genetic diversity was relatively high with samples 20 m apart representing genetically distinct individuals (Fatemi and Gross, unpublished data). It remains unknown what factors influence the mode of reproduction in different populations in Australia but it is likely that the type of flood event and propagule availability are important. Previous field studies have found that seedling recruitment was the predominant reproductive mode (approx. 70%) following winter flooding, in single site studies (Macdonald 2008; Price et al. 2010a). We examine the density of recruits, dispersal distance and direction from adult plants, and mode of reproduction in a semi-arid floodplain. The study area J. N. Price M. J. Macdonald C. L. Gross I. H. Simpson Ecosystem Management, University of New England, Armidale, NSW 2351, Australia

Microsatellite loci in the carcinogenic liver fluke, Opisthorchis viverrini and their application as population genetic markers

Opisthorchis viverrini is a carcinogenic foodborne trematode endemic in Southeast Asia especially in Thailand and the Lao People's Democratic Republic. Opisthorchiasis causes hepatobiliary diseases and cholangiocarcinoma (bile duct cancer). Currently there is substantial evidence on genetic variation of O. viverrini but the information on population genetic structure is lacking. Because microsatellite DNA of this parasite is not available, we for the first time isolated and utilized microsatellite DNA as genetic markers to examine genetic diversity and the population structure of O. viverrini. Partial genomic DNA libraries were constructed by conventional and enrichment methods which yielded microsatellite-containing clones of 0.18–0.25% and 16.84%, respectively. Within 41 microsatellite loci isolated 36.59% were perfect, 60.98% were interrupted and 2.44% were compound microsatellites. The CA repetitions were the most frequent, followed by GT and CAT. Primers specific to the flanking regions of 12 microsatellite loci were developed to genotype 150 O. viverrini individuals from geographical localities in Thailand and Lao PDR. Allele numbers per locus ranged from 2 to 15, with the mean expected heterozygosity of 0.03–0.66. Analyses of O. viverrini from 5 localities revealed a high level of genetic diversity and had significant deviation from Hardy–Weinberg equilibrium. Significant heterozygote deficiency as well as heterozygote excess was detected across all localities indicating the possibility of selfing (inbreeding) as a predominant reproductive mode. Significant genetic differentiation ( F ST) was also detected between worms from different localities with varying levels of genetic heterogeneity. We discuss our results in terms of what these novel microsatellite markers reveal about the epidemiology and transmission dynamics of this medically important parasite, both in terms of the current study and their potential for future comprehensive population genetic studies O. viverrini sensu lato in Southeast Asia.

Mutation load and rapid adaptation favour outcrossing over self-fertilization

The tendency of organisms to reproduce by cross-fertilization despite numerous disadvantages relative to self-fertilization is one of the oldest puzzles in evolutionary biology. For many species, the primary obstacle to the evolution of outcrossing is the cost of production of males, individuals that do not directly contribute offspring and thus diminish the long-term reproductive output of a lineage. Self-fertilizing ('selfing') organisms do not incur the cost of males and therefore should possess at least a twofold numerical advantage over most outcrossing organisms. Two competing explanations for the widespread prevalence of outcrossing in nature despite this inherent disadvantage are the avoidance of inbreeding depression generated by selfing and the ability of outcrossing populations to adapt more rapidly to environmental change. Here we show that outcrossing is favoured in populations of Caenorhabditis elegans subject to experimental evolution both under conditions of increased mutation rate and during adaptation to a novel environment. In general, fitness increased with increasing rates of outcrossing. Thus, each of the standard explanations for the maintenance of outcrossing are correct, and it is likely that outcrossing is the predominant mode of reproduction in most species because it is favoured under ecological conditions that are ubiquitous in natural environments.

Local genetic structure in a North American epiphytic lichen, Ramalina menziesii (Ramalinaceae)

Epiphytic lichens possess unique life history traits that can have conflicting effects on genetic structure: symbiotic mutualism between a fungus with its algal or cyanobacterial photobiont, association with a host plant, and ability to reproduce sexually and asexually. Our study species, Ramalina menziesii, has small ascospores that can facilitate long-distance gene movement, and it is capable of clonal reproduction. The goals of this study are to test whether different haplotypes were differentially distributed across host plant species, to look for evidence of asexual vs. sexual reproduction, and to assess the local genetic structure of the population. We sampled individuals from multiple trees of three oak species in four lichen subpopulations within a savanna ecosystem. Using DNA sequence data from four fungal nuclear loci, we found no tendency for host specialization. Alleles were randomly distributed across subpopulations. The frequency of multilocus genotypes was consistent with a randomly mating population. Sexual reproduction involving relichenization appeared to be the predominant mode of reproduction of R. menziesii at this study site. We found no significant local genetic structure suggesting widespread gene flow at the local scale. The genetic structure of this lichen is comparable to that of widely distributed epiphytic plants.

The Evolutionary Maintenance of Sexual Reproduction: Evidence from the Ecological Distribution of Asexual Reproduction in Clonal Plants

In theory, females that reproduce asexually should enjoy a twofold advantage in fitness over sexual females, yet sex remains the predominant mode of reproduction in virtually all eukaryotes. The evolutionary maintenance of sex is especially puzzling in clonal plants because the transition from sexual to exclusively asexual reproduction is an ever‐present possibility in these species. In this article, I use published data on the genotypic diversity of populations of clonal plants to test five hypotheses about the ecological situations that limit or favor clonal reproduction in vascular plants. The data were drawn from 248 studies covering 69,000 individuals in >2000 populations of 218 species in 74 plant families. The tests showed the following: (1) the frequency of clonality increases with population age, indicating that clonal reproduction is limited by disturbance; (2) clonal reproduction is limited by dispersal; clones are more frequent in aquatic and apomictic species in which the dispersal of clonally produced propagules is less limiting; (3) clones are more frequent in populations of rare or endangered species; (4) populations of alien plants have higher frequencies of clonality; and (5) clones are more frequent at the edges of species’ geographical ranges. Thus, it appears that the ultimately successful clonal plant would be a rare, aquatic, alien apomict living in an undisturbed, geographically marginal habitat. Since this combination of circumstances is so restrictive, it is perhaps better regarded as a sign of sexual failure than as a recipe for clonal success.