Research Items (140)
Multiple infections (co‐occurrence of multiple pathogen genotypes within an individual host) can have important impacts on diseases. Relatedness among pathogens can affect the likelihood of multiple infections and their consequences through kin selection. Previous studies on the castrating anther‐smut fungus Microbotryum lychnidis‐dioicae have shown that multiple infections occur in its host plant Silene latifolia. Relatedness was high among fungal genotypes within plants, which could result from competitive exclusion between unrelated fungal genotypes, from population structure or from interactions between plant and fungal genotypes for infection ability. Here, we aimed at disentangling these hypotheses using M. saponariae and its host Saponaria officinalis, both experimentally tractable for these questions. By analysing populations using microsatellite markers, we also found frequent occurrence of multiple infections and high relatedness among strains within host plants. Infections resulting from experimental inoculations in the greenhouse also revealed high relatedness among strains coinfecting host plants, even in clonally‐replicated plant genotypes, indicating that high relatedness within plants did not result merely from plant x fungus interactions or population structure. Furthermore, hyphal growth in vitro was affected by the presence of a competitor growing nearby and by its genetic similarity, although this latter effect was strain‐dependent. Altogether, our results support the hypothesis that relatedness‐dependent competitive exclusion occurs in Microbotryum fungi within plants. These micro‐organisms can thus respond to competitors and to their level of relatedness. This article is protected by copyright. All rights reserved.
Some bacteria produce and perceive quorum-sensing (QS) signals that coordinate several behaviours, including the costly processes that are exoenzyme production and plasmid transfer. In the case of plasmid transfer, the emergence of QS signal-altered invaders and their policing are poorly documented. In Agrobacterium tumefaciens, the virulence Ti-plasmid encodes both synthesis and sensing of QS-signals, which promote its transfer from a donor to a recipient cell. Here, we reported that QS-altered A. tumefaciens mutants arose during experimental evolution. All showed improved growth compared to their ancestor. Genome sequencing revealed that, though some had lost the Ti-plasmid, most were defective for QS-signal synthesis and Ti-plasmid conjugation (traR mutations) and one exhibited a QS-signal exploitation behaviour, using signal produced by other cells to enhance its own Ti-plasmid transfer. We explored mechanisms that can limit this QS-hijacking. We showed that the A. tumefaciens capacity to inactivate QS-signals by expressing QS-degrading enzyme could attenuate dissemination of the QS signal-negative Ti-plasmids. This work shows that enzymatic QS-disruption whether encoded by the QS-producing Ti-plasmid itself, by a companion plasmid in the same donor cells, or by one in the recipient cells, in all cases can serve as a mechanism for controlling QS exploitation by QS signal-negative mutants.
Background Anther-smut fungi belonging to the genus Microbotryum sterilize their host plants by aborting ovaries and replacing pollen by fungal spores. Sibling Microbotryum species are highly specialized on their host plants and they have been widely used as models for studies of ecology and evolution of plant pathogenic fungi. However, most studies have focused, so far, on M. lychnidis-dioicae that parasitizes the white campion Silene latifolia. Microbotryum saponariae, parasitizing mainly Saponaria officinalis, is an interesting anther-smut fungus, since it belongs to a tetrapolar lineage (i.e., with two independently segregating mating-type loci), while most of the anther-smut Microbotryum fungi are bipolar (i.e., with a single mating-type locus). Saponaria officinalis is a widespread long-lived perennial plant species with multiple flowering stems, which makes its anther-smut pathogen a good model for studying phylogeography and within-host multiple infections. Principal Findings Here, based on a generated genome sequence of M. saponariae we developed 6 multiplexes with a total of 22 polymorphic microsatellite markers using an inexpensive and efficient method. We scored these markers in fungal individuals collected from 97 populations across Europe, and found that the number of their alleles ranged from 2 to 11, and their expected heterozygosity from 0.01 to 0.58. Cross-species amplification was examined using nine other Microbotryum species parasitizing hosts belonging to Silene, Dianthus and Knautia genera. All loci were successfully amplified in at least two other Microbotryum species. Significance These newly developed markers will provide insights into the population genetic structure and the occurrence of within-host multiple infections of M. saponariae. In addition, the draft genome of M. saponariae, as well as one of the described markers will be useful resources for studying the evolution of the breeding systems in the genus Microbotryum and the evolution of specialization onto different plant species.
Capillary electrophoresis chromatograms with peaks of heterozygous individuals of Microbotryum saponariae at the polymorphic microsatellite loci 102 (Msap_102). Heterozygous individuals have the following allelic profile: a) 173 and 187; b) 173 and 196; c) 173 and 184. (PDF)
Information on the Microbotryum individuals sampled from different host plants across Europe and analysed in this study. Individual fungal DNA was extracted using Chelex protocol (Biorad, USA) and was used to test the polymorphism of the 22 microsatellite markers in M. saponariae and in the other Microbotryum species used for cross-species amplification. Symbols (*) refers to DNA samples that were also extracted with the Nucleospin Soil kit (Macherey-Nagel, Germany) and were pooled as a template DNA for downstream applications in the first screening of markers polymorphism. (PDF)
- Nov 2016
The facultative hemiparasitic plant Rhamphicarpa fistulosa (Orobanchaceae) thrives in seasonally wet soils in sub-Saharan Africa, mainly in marginal lowland rice growing environments where weeds are already a major constraint for rice production. Because lowland rice production is increasing in tropical Africa, it is important to ascertain the influence of R. fistulosa on weed plant communities in these rice-growing habitats. We investigated weed plant community richness and composition at four different levels of R. fistulosa infestation across two years of surveys from lowland rice fields in northern Togo (West Africa). Despite a lack of significant differences in community richness among sites with different R. fistulosa infestation levels, there were significant differences in community composition, both when estimated from presence-absence data and from relative abundance data, after controlling statistically for geographic proximity among sites. Rhamphicarpa fistulosa infestation, therefore, may influence the competitive balance between rice and its weeds and shape weed community structure. However, experimental studies are required to elucidate the weed host range of R. fistulosa and the direct and indirect effects of this hemiparasite in rice fields in order to predict its net impact on rice and its weed species.
We investigated the molecular and ecological mechanisms involved in niche expansion, or generalism, versus specialization in sympatric plant pathogens. Nopaline-type and octopine-type Agrobacterium tumefaciens engineer distinct niches in their plant hosts that provide different nutrients: nopaline or octopine, respectively. Previous studies revealed that nopaline-type pathogens may expand their niche to also assimilate octopine in the presence of nopaline, but consequences of this phenomenon on pathogen dynamics in planta were not known. Here, we provided molecular insight into how the transport protein NocT can bind octopine as well as nopaline, contributing to niche expansion. We further showed that despite the ability for niche expansion, nopaline-type pathogens had no competitive advantage over octopine-type pathogens in co-infected plants. We also demonstrated that a single nucleotide polymorphism in the nocR gene was sufficient to allow octopine assimilation by nopaline-type strains even in absence of nopaline. The evolved nocR bacteria had higher fitness than their ancestor in octopine-rich transgenic plants but lower fitness in tumors induced by octopine-type pathogens. Overall, this work elucidates the specialization of A. tumefaciens to particular opine niches and explains why generalists do not always spread despite the advantage associated with broader nutritional niches.The ISME Journal advance online publication, 1 November 2016; doi:10.1038/ismej.2016.137.
Crop-to-wild gene flow can reduce the fitness and genetic integrity of wild species. Malus sylvestris, the European crabapple fruit tree, in particular is threatened by the disappearance of its habitat and by gene flow from its domesticated relative, Malus domestica. With the aims of evaluating threats for M. sylvestris and of formulating recommendations for its conservation, we studied here, using microsatellite markers and growth experiments: i) hybridization rates in seeds and trees from a French forest and in seeds used for replanting crabapples in agrosystems and in forests, ii) the impact of the level of M. domestica ancestry on individual fitness tree, and iii) pollen dispersal abilities in relationship with crop-to-wild gene flow. We found substantial contemporary crop-to-wild gene flow in crabapple tree populations and superior fitness of hybrids compared to wild seeds and seedlings. Using paternity analyses, we showed that pollen dispersal could occur up to 4 km and decreased with tree density. The seed network furnishing the wild apple reintroduction agroforestry programs was found to suffer from poor genetic diversity, introgressions, and species misidentification. Overall, our findings indicate supported threats for the European wild apple steering us to provide precise recommendations for its conservation. This article is protected by copyright. All rights reserved.
- May 2016
Nuclear disasters at Chernobyl and Fukushima provide examples of effects of acute ionizing radiation on mutations that can affect the fitness and distribution of species. Here, we investigated the prevalence of Microbotryum lychnidis-dioicae, a pollinator-transmitted fungal pathogen of plants causing anther-smut disease in Chernobyl, its viability, fertility and karyotype variation, and the accumulation of non-synonymous mutations in its genome. We collected diseased flowers of Silene latifolia from locations ranging by more than two orders of magnitude in background radiation, from 0.05 to 21.03 μSv/h. Disease prevalence decreased significantly with increasing radiation level, possibly due to lower pollinator abundance and altered pollinator behaviour. Viability and fertility, measured as the budding rate of haploid sporidia following meiosis from the diploid teliospores, did not vary with increasing radiation levels and neither did karyotype overall structure and level of chromosomal size heterozygosity. We sequenced the genomes of twelve samples from Chernobyl and of four samples collected from uncontaminated areas and analysed alignments of 6,068 predicted genes, corresponding to 1.04x10(7) base pairs. We found no dose-dependent differences in substitution rates (neither dN, dS, nor dN/dS). Thus, we found no significant evidence of increased deleterious mutation rates at higher levels of background radiation in this plant pathogen. We even found lower levels of non-synonymous substitution rates in contaminated areas compared to control regions, suggesting that purifying selection was stronger in contaminated than uncontaminated areas. We briefly discuss the possibilities for a mechanistic basis of radio-resistance in this non-melanised fungus. This article is protected by copyright. All rights reserved.
- Oct 2015
The evolutionary equilibrium hypothesis was proposed to explain variation in egg rejection rates among individual hosts (intra- and interspecific) of avian brood parasites. Hosts may sometimes mistakenly reject own eggs when they are not parasitized (i.e. make recognition errors). Such errors would incur fitness costs and could counter the evolution of host defences driven by costs of parasitism (i.e. creating equilibrium between acceptors and rejecters within particular host populations). In the present study, we report the disappearance of host eggs from nonparasitized nests in populations of seven actual and potential hosts of the common cuckoo Cuculus canorus. Based on these data, we calculate the magnitude of the balancing parasitism rate provided that all eggs lost are a result of recognition errors. Importantly, because eggs are known to disappear from nests for reasons other than erroneous host rejection, our data represent the maximum estimates of such costs. Nonetheless, the disappearance of eggs was a rare event and therefore incurred low costs compared to the high costs of parasitism. Hence, costs as a result of recognition errors are probably of minor importance with respect to opposing selective pressure for the evolution of egg rejection in these hosts. We cannot exclude the possibility that low or intermediate egg rejection rates in some host populations may be caused by spatiotemporal variation in the occurrence of parasitism and gene flow, creating a variable influence of opposing costs as a result of recognition errors and the costs of parasitism.
Résumé Rhamphicarpa fistulosa (Hochst.) Benth. (Scrophulariaceae) est une plante adventice hémiparasite épirhize annuelle rencontrée dans les bas-fonds et les plaines alluviales en Afrique tropicale. Au nord du Togo, dans la région des Savanes, elle a été signalée comme la contrainte biotique majeure pour la riziculture de bas-fonds. Une prospection a été réalisée en 2012 pour déterminer l'étendue de son infestation. Une enquête basée sur un questionnaire semi-ouvert a été effectuée auprès des paysans des parcelles visitées. Rhamphicarpa fistulosa a été retrouvée dans 80 % des 33 bas-fonds prospectés. La période d'abondance s'étend d'août à septembre, où les plus forts niveaux d'infestation (90 %) ont été observés sur des sols sableux et argileux. Lorsque R. fistulosa est présente, 94 autres plantes adventices issues de 24 familles botaniques sont également recensées. Comme méthode de lutte, les paysans utilisent le sarclage et l'épandage d'engrais de synthèse, mais toujours à des taux inférieurs aux recommandations nationales. L'abandon des parcelles à la suite d'une forte infestation a été constaté ainsi que d'autres dégâts (réduction du développement des plants de riz et leur jaunissement). Mots clés : agroécologie ; relation hôte-parasite ; plante parasite ; riz.
The level of parasite virulence, i.e., the decrease in host's fitness due to a pathogen, is expected to depend on several parameters, such as the type of the disease (e.g., castrating or host-killing) and the prevalence of multiple infections. Although these parameters have been extensively studied theoretically, few empirical data are available to validate theoretical predictions. Using the anther smut castrating disease on Silene latifolia caused by Microbotryum lychnidis-dioicae, we studied the dynamics of multiple infections and of different components of virulence (host death, non-recovery and percentage of castrated stems) during the entire lifespan of the host in an experimental population. We monitored the number of fungal genotypes within plants and their relatedness across five years, using microsatellite markers, as well as the rates of recovery and host death in the population. The mean relatedness among genotypes within plants remained at a high level throughout the entire host lifespan despite the dynamics of the disease, with recurrent new infections. Recovery was lower for plants with multiple infections compared to plants infected by a single genotype. As expected for castrating parasites, M. lychnidis-dioicae did not increase host mortality. Mortality varied across years but was generally lower for plants that had been diseased the preceding year. This is one of the few studies to have empirically verified theoretical expectations for castrating parasites, and to show particularly i) that castrated hosts live longer, suggesting that parasites can redirect resources normally used in reproduction to increase host lifespan, lengthening their transmission phase, and ii) that multiple infections increase virulence, here in terms of non-recovery and host castration.
Rhamphicarpa fistulosa (Hochst.) Benth. is an annual facultative root-parasitic weed found in lowlands and floodplains in tropical Africa. It has been reported as the major constraint to rice cultivation in the savannah region of Togo. A survey was conducted in 2012 to access its distribution and to collect information on farmers’ knowledge of its control. Rhamphicarpa fistulosa was found in 80% of the 33 visited lowlands. The period of abundance is in August-September when the highest infection (90%) was found on sandy and clay soils. When the parasite was present, 94 other weeds belonging to 24 botanical families were recorded. As management method, farmers use weeding and chemical fertilizers application but always at lower input rates than national standards. The abandonment of plots is mainly due to a heavy infection, other reported damages are the reduction of rice plants growth and plant yellowing.
- Dec 2012
Although plant resistance to aggressors has been well described, there is still little knowledge about the mechanisms underlying their tolerance to pathogens. Tolerance often appears to be mediated by changes in life history traits, shifting host resource investment from growth to reproduction, but whether host phenotype modifications induced after attack are adaptive is not always clear. Here, we investigated the details of the impact of Hyaloperonospora arabidopsidis infection on several biomass, phenology and architectural traits of Arabidopsis thaliana, for three pathogen genotypes and three host plant genotypes that have been shown previously to differ greatly in fecundity and tolerance to infection. We found that, although host genotype explains most of the variance in life history traits, these three lines differ critically in their response to infection, with delays and biomass losses at bolting, together with changes in inflorescence architecture, observed at one extreme host line, and an advantage at bolting for infected plants and no inflorescence alteration for the other. These results suggest that the differences in tolerance observed previously in this pathosystem do not involve plasticity in inflorescence architecture, but may arise from induced changes at the vegetative stage, before plant transition to reproduction.
- May 2012
In avian brood parasitism, egg phenotype plays a key role for both host and parasite reproduction. Several parrotbill species of the genus Paradoxornis are parasitized by the common cuckoo Cuculus canorus, and clear polymorphism in egg phenotype is observed. In this article, we develop a population genetics model in order to identify the key parameters that control the maintenance of egg polymorphism. The model analyses show that egg polymorphism can be maintained either statically as an equilibrium or dynamically with frequency oscillations depending on the sensitivity of the host against unlike eggs and how the parasite targets host nests with specific egg phenotypes. On the basis of the model, we discuss egg polymorphism observed in parrotbills and other host species parasitized by the cuckoo. We suggest the possibility that frequencies of egg phenotypes oscillate and we appeal for monitoring of cuckoo–host interactions over a large spatiotemporal scale.
Avian brood parasites impose strong selection on their hosts leading to the evolution of antiparasite defenses like egg recognition and rejection. Discordance and template-based cognitive mechanisms may form the base for egg recognition by hosts. For discordance, hosts recognize eggs that constitute the minority in a clutch as alien, whereas in template-based recognition, hosts recognize eggs as alien when they do not match a template that can be innate or learnt. Template-based recognition by learning can be compromised in host species with polymorphic egg color like Paradoxornis parrotbills, hosts of the common cuckoo Cuculus canorus, because a male that learns an egg color in his first breeding attempt can subsequently mate with females having different colors and therefore reject his own eggs. We present a simple conceptual model to understand how an asymmetry in sex roles of care for eggs and egg polymorphism influence the evolution of egg recognition by hosts. We derive host reproductive success in the presence of variation in egg phenotype for both host and parasite. Our model shows that male recognition by learning is disadvantageous unless the host has monomorphic eggs. We suggest that interclutch variation in egg phenotype is the key to understanding the evolution of egg recognition and the sex involved.
Avian brood parasites and their hosts are involved in complex offence-defense coevolutionary arms races. The most common pair of reciprocal adaptations in these systems is egg discrimination by hosts and egg mimicry by parasites. As mimicry improves, more advanced host adaptations evolve such as decreased intra- and increased interclutch variation in egg appearance to facilitate detection of parasitic eggs. As interclutch variation increases, parasites able to choose hosts matching best their own egg phenotype should be selected, but this requires that parasites know their own egg phenotype and select host nests correspondingly. We compared egg mimicry of common cuckoo Cuculus canorus eggs in naturally parasitized marsh warbler Acrocephalus palustris nests and their nearest unparasitized conspecific neighbors having similar laying dates and nest-site characteristics. Modeling of avian vision and image analyses revealed no evidence that cuckoos parasitize nests where their eggs better match the host eggs. Cuckoo eggs were as good mimics, in terms of background and spot color, background luminance, spotting pattern and egg size, of host eggs in the nests actually exploited as those in the neighboring unparasitized nests. We reviewed the evidence for brood parasites selecting better-matching host egg phenotypes from several relevant studies and argue that such selection probably cannot exist in host-parasite systems where host interclutch variation is continuous and overall low or moderate. To date there is also no evidence that parasites prefer certain egg phenotypes in systems where it should be most advantageous, i.e., when both hosts and parasites lay polymorphic eggs. Hence, the existence of an ability to select host nests to maximize mimicry by brood parasites appears unlikely, but this possibility should be further explored in cuckoo-host systems where the host has evolved discrete egg phenotypes.
Although parasites and their hosts often coexist in a set of environmentally differentiated populations connected by gene flow, few empirical studies have considered a role of environmental variation in shaping correlations between traits of hosts and parasites. Here, we studied for the first time the association between the frequency of adaptive parasitic common cuckoo Cuculus canorus phenotypes in terms of egg matching and level of defences exhibited by its reed warbler Acrocephalus scirpaceus hosts across seven geographically distant populations in Europe. We also explored the influence of spring climatic conditions experienced by cuckoos and hosts on cuckoo-host egg matching. We found that between-population differences in host defences against cuckoos (i.e. rejection rate) covaried with between-population differences in degree of matching. Between-population differences in host egg phenotype were associated with between-population differences in parasitism rate and spring climatic conditions, but not with host level of defences. Between-population differences in cuckoo egg phenotype covaried with between-population differences in host defences and spring climatic conditions. However, differences in host defences still explained differences in mimicry once differences in climatic conditions were controlled, suggesting that selection exerted by host defences must be strong relative to selection imposed by climatic factors on egg phenotypes.
An official journal of the Genetics Society, Heredity publishes high-quality articles describing original research and theoretical insights in all areas of genetics. Research papers are complimented by News & Commentary articles and reviews, keeping researchers and students abreast of hot topics in the field.
Gynodioecy, the coexistence of female and hermaphrodite plants within a species, is often under nuclear-cytoplasmic sex determination, involving cytoplasmic male sterility (CMS) genes and nuclear restorers. A good knowledge of CMS and restorer polymorphism is essential for understanding the evolution and maintenance of gynodioecy, but reciprocal crossing studies remain scarce. Although mitochondrial diversity has been studied in a few gynodioecious species, the relationship between mitotype diversity and CMS status is poorly known. From a French sample of Silene nutans, a gynodioecious species whose sex determination remains unknown, we chose the four most divergent mitotypes that we had sampled at the cytochrome b gene and tested by reciprocal crosses whether they carry distinct CMS genes. We show that gynodioecy in S. nutans is under nuclear-cytoplasmic control, with at least two different CMSs and up to four restorers with epistatic interactions. Female occurrence and frequency were highly dependent on the mitotype, suggesting that the level of restoration varies greatly among CMSs. Two of the mitotypes, which have broad geographic distributions, represent different CMSs and are very unequally restored. We discuss the dynamics of gynodioecy at the large-scale meta-population level.
The common ragweed (Ambrosia artemisiifolia L.; Asteraceae) is a North American native that is invading Eurasia. Besides its economic impact on crop yield, it presents a major health problem because of its highly allergenic pollen. The plant was imported inadvertently to Europe in the eighteenth century and has become invasive in several countries. By analyzing French and North American populations, it was previously shown that French populations were best described as a mixture of native sources and that range expansion in France probably involved sequential bottlenecks. Here, our aim was to determine whether Eastern European populations of A. artemisiifolia originated from the previously established French populations or from independent trans-Atlantic colonization events. We used nuclear microsatellite markers to elucidate the relationships among populations from Eastern and Western Europe in relation to populations from a broad survey across the native North American range. We found that A. artemisiifolia from Eastern Europe did not originate from the earlier established French populations but rather represents multiple independent introductions from other sources, or introductions from a not yet identified highly diverse native population. Eastern European populations show comparable amounts of genetic variability as do previously characterized French and North American populations, but analyses of population structure clearly distinguish the two European groups. This suggests separate introductions in Eastern and Western Europe as well as divergent sources for these two invasions, possibly as a result of distinct rules for trade and exchange for Eastern Europe during most of the twentieth century. KeywordsAllergenic plant–Biological invasion–Invasive species–Multiple introductions–Population structure
- Apr 2011
Understanding the mechanisms responsible for divergence and specialization of pathogens on different hosts is of fundamental importance, especially in the context of the emergence of new diseases via host shifts. Temporal isolation has been reported in a few plants and parasites, but is probably one of the least studied speciation processes. We studied whether temporal isolation could be responsible for the maintenance of genetic differentiation among sympatric populations of Ampelomyces, widespread intracellular mycoparasites of powdery mildew fungi, themselves plant pathogens. The timing of transmission of Ampelomyces depends on the life cycles of the powdery mildew species they parasitize. Internal transcribed spacer sequences and microsatellite markers showed that Ampelomyces populations found in apple powdery mildew (Podosphaera leucotricha) were genetically highly differentiated from other Ampelomyces populations sampled from several other powdery mildew species across Europe, infecting plant hosts other than apple. While P. leucotricha starts its life cycle early in spring, and the main apple powdery mildew epidemics occur before summer, the fungal hosts of the other Ampelomyces cause epidemics mainly in summer and autumn. When two powdery mildew species were experimentally exposed to Ampelomyces strains naturally occurring in P. leucotricha in spring, and to strains naturally present in other mycohost species in autumn, cross-infections always occurred. Thus, the host-related genetic differentiation in Ampelomyces cannot be explained by narrow physiological specialization, because Ampelomyces were able to infect powdery mildew species they were unlikely to have encountered in nature, but instead appears to result from temporal isolation.
Relationship between geographic and genetic distances between all pairs of populations based on nDNA loci. A) In North America (P = 0.002). B) In Europe (P = 0.581). The geographic distance was expressed as the log10 of interpopulation distance in km; the genetic distance was expressed as FST/(1-FST). (TIF)
CpDNA haplotypic composition of the studied populations. Populations are grouped into three spatial groups (North America, Europe and non-European invasive populations) and roughly ordered from West to East. Haplotypes are ordered from the most frequent to the least frequent. Haplotypes that are in bold are private to one population. (XLS)
Pairwise FST indices among all pairs of populations, estimated at nDNA loci. Populations are grouped into three spatial groups (North America, Europe and non-European invasive populations) and roughly ordered from West to East. FST estimates that are highlighted in grey correspond to non-significant exact tests of differentiation. (XLS)
Results of the assignment test of invasive populations to North American populations, based on nDNA loci. The number of individuals assigned to each North American population is given, either considering all individuals or only individuals with assignment probabilities above 0.5. (XLS)
Ambrosia artemisiifolia is a North American native that has become one of the most problematic invasive plants in Europe and Asia. We studied its worldwide population genetic structure, using both nuclear and chloroplast microsatellite markers and an unprecedented large population sampling. Our goals were (i) to identify the sources of the invasive populations; (ii) to assess whether all invasive populations were founded by multiple introductions, as previously found in France; (iii) to examine how the introductions have affected the amount and structure of genetic variation in Europe; (iv) to document how the colonization of Europe proceeded; (v) to check whether populations exhibit significant heterozygote deficiencies, as previously observed. We found evidence for multiple introductions of A. artemisiifolia, within regions but also within populations in most parts of its invasive range, leading to high levels of diversity. In Europe, introductions probably stem from two different regions of the native area: populations established in Central Europe appear to have originated from eastern North America, and Eastern European populations from more western North America. This may result from differential commercial exchanges between these geographic regions. Our results indicate that the expansion in Europe mostly occurred through long-distance dispersal, explaining the absence of isolation by distance and the weak influence of geography on the genetic structure in this area in contrast to the native range. Last, we detected significant heterozygote deficiencies in most populations. This may be explained by partial selfing, biparental inbreeding and/or a Wahlund effect and further investigation is warranted. This insight into the sources and pathways of common ragweed expansion may help to better understand its invasion success and provides baseline data for future studies on the evolutionary processes involved during range expansion in novel environments.
Parasites require synchrony with their hosts so if host timing changes with climate change, some parasites may decline and eventually go extinct. Residents and short-distance migrant hosts of the brood parasitic common cuckoo, Cuculus canorus, have advanced their phenology in response to climate change more than long-distance migrants, including the cuckoo itself. Because different parts of Europe show different degrees of climate change, we predicted that use of residents or short-distance migrants as hosts should have declined in areas with greater increase in spring temperature. Comparing relative frequency of parasitism of the two host categories in 23 European countries before and after 1990, when spring temperatures in many areas had started to increase, we found that relative parasitism of residents and short-distance migrants decreased. This change in host use was positively related to increase in spring temperature, consistent with the prediction that relative change in phenology for different migrant classes drives host-use patterns. These findings are consistent with the hypothesis that climate change affects the relative abundance of different host races of the common cuckoo.
- Feb 2011
Nectar production and concentration were measured on male and female plants of six experimental lines of Silene latifolia (Caryophyllaceae) that differ in disease resistance. Differences between the sexes and among the lines were found for nectar production, concentration, and total sugar production during the first 24 h of anthesis. Females produced more nectar of lower concentration than did males, and males secreted more sugar than did females during the first day of anthesis. However, nectar traits of males and females resembled one another within a line, suggesting a genetic correlation between the sexes. Further, the additive genetic basis for this trait appears weak. In S. latifolia repeatabilities for nectar traits were low and differed between females and males, so the heritabilities must be extremely low. Groups of plant lines that are "susceptible" and "resistant" to the fungal pathogen Microbotryum violaceum did not differ in nectar parameters. Therefore resistance to this pollinator-borne disease does not appear to influence nectar production. Key words: Microbotryum (= Ustilago), plant – pathogen interactions, pollinator reward, anther-smut disease, dioecy, floral nectar.
- Jan 2011
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- Dec 2010
Critical determinants of the optimum level of virulence in pathogens include the presence of competitors (i.e., multiple infections), their relatedness, and the effect of competitors on pathogen growth and disease development. Empirical data regarding the existence of competitive interactions and their impact on virulence remain very limited compared to theoretical studies. Here, we followed an experimental population of the model fungal pathogen Microbotryum lychnidis-dioicae on its caryophyllaceous host Silene latifolia. Our analysis revealed conditional responses by the pathogen to the presence of competitors, which was dependent upon the relatedness of pathogens within hosts. Overall, virulence was increased in cases of multiple infections as compared to single infections: both spore production and degree of plant sterilization were higher under multiple infections. The pathogen indeed increased its growth and reproductive rate when competitors were present within the same plant. Microbotryum also appeared able to interfere with competitors, reducing their ability to colonize the host, and this effect was smaller between closer relatives. Our results thus help to elucidate the myriad of theoretical considerations on the evolution of virulence by providing experimental results with a well-studied disease of wild plant populations.
The obligate avian brood parasitic common cuckoo Cuculus canorus comprises different strains of females that specialize on particular host species by laying eggs of a constant type that often mimics those of the host. Whether cuckoos are locally adapted for mimicking populations of the hosts on which they are specialized has never been investigated. In this study, we first explored the possibility of local adaptation in cuckoo egg mimicry over a geographical mosaic of selection exerted by one of its main European hosts, the reed warbler Acrocephalus scirpaceus. Secondly, we investigated whether cuckoos inhabiting reed warbler populations with a broad number of alternative suitable hosts at hand were less locally adapted. Cuckoo eggs showed different degrees of mimicry to different reed warbler populations. However, cuckoo eggs did not match the egg phenotypes of their local host population better than eggs of other host populations, indicating that cuckoos were not locally adapted for mimicry on reed warblers. Interestingly, cuckoos exploiting reed warblers in populations with a relatively larger number of co-occurring cuckoo gentes showed lower than average levels of local adaptation in egg volume. Our results suggest that cuckoo local adaptation might be prevented when different cuckoo populations exploit more or fewer different host species, with gene flow or frequent host switches breaking down local adaptation where many host races co-occur.
Generalist parasites regularly evolve host-specific races that each specialize on one particular host species. Many host-specific races originate from geographically structured populations where local adaptations to different host species drive the differentiation of distinct races. However, in sympatric populations where several host races coexist, gene flow could potentially disrupt such host-specific adaptations. Here, we analyse genetic differentiation among three sympatrically breeding host races of the brood-parasitic common cuckoo, Cuculus canorus. In this species, host-specific adaptations are assumed to be controlled by females only, possibly via the female-specific W-chromosome, thereby avoiding that gene flow via males disrupts local adaptations. Although males were more likely to have offspring in two different host species (43% versus 7%), they did not have significantly more descendants being raised outside their putative foster species than females (9% versus 2%). We found significant genetic differentiation for both biparentally inherited microsatellite DNA markers and maternally inherited mitochondrial DNA markers. To our knowledge, this is the first study that finds significant genetic differentiation in biparentally inherited markers among cuckoo host-specific races. Our results imply that males also may contribute to the evolution and maintenance of the different races, and hence that the genes responsible for egg phenotype may be found on autosomal chromosomes rather than the female-specific W-chromosome as previously assumed.
- Mar 2010
The brood parasitic common cuckoo Cuculus canorus consists of gentes, which typically parasitize only a single host species whose eggs they often mimic. Where multiple cuckoo gentes co-exist in sympatry, we may expect variable but generally poorer mimicry because of host switches or inter-gens gene flow via males if these also contribute to egg phenotypes. Here, we investigated egg trait differentiation and mimicry in three cuckoo gentes parasitizing great reed warblers Acrocephalus arundinaceus, marsh warblers Acrocephalus palustris and corn buntings Miliaria calandra breeding in close sympatry in partially overlapping habitat types. The three cuckoo gentes showed a remarkable degree of mimicry to their three host species in some but not all egg features, including egg size, a hitherto largely ignored feature of egg mimicry. Egg phenotype matching for both background and spot colours as well as for egg size has been maintained in close sympatry despite the possibility for gene flow.
In order to produce seeds, animal-pollinated plants must flower synchronously with and be attractive to their pollinators while avoiding antagonists. Here, we explore temporal and inter-individual patterns in pollination and pre-dispersal seed predation of Dianthus sylvestris by Hadena moths, within and among three sex morphs. We scored plants that started flowering at different periods in 2001 and 2003 and found that fruit set decreased and predation rates increased over one season, and most of the other season, granting a female reproductive advantage to early flowering plants, though, we found no morph-specific temporal patterns. Female plants set more fruits, and more of their fruits escaped predation in one year, but this did not grant them a reproductive advantage since they produce fewer flowers per plant than the other morphs. Instead, mixed plants showed a clear female reproductive advantage. We also examined predation types by Hadena and seed production in attacked and intact fruits of individually marked flowers. Though female Hadena moths laid eggs preferentially into perfect flowers, flower sexes suffered similar predation by itinerant caterpillars. Attacked fruits contained fewer and lighter seeds than un-attacked ones. We conclude that pre-dispersal seed predation by Hadena may select on flowering onset of this sexually polymorphic species.
Predominantly outcrossing plant species are expected to accumulate recessive deleterious mutations, which can be purged when in a homozygous state following selfing. Individuals may vary in their genetic load because of different selfing histories, which could lead to differences in inbreeding depression among families. Lineage-dependent inbreeding depression can appear in gynodioecious species if obligatory outcrossed females are more likely to produce female offspring and if partially selfing hermaphrodites are more likely to produce hermaphrodites. We investigated inbreeding depression at the zygote, seed, and germination stages in the gynomonoecious-gynodioecious Dianthus sylvestris, including pure-sexed plants and a mixed morph. We performed hand-pollinations on 56 plants, belonging to the three morphs, each receiving 2-3 cross treatments (out-, sib- and self-pollination) on multiple flowers. Effects of cross treatments varied among stages and influenced seed provisioning, with sibling competition mainly occurring within outcrossed fruits. We found significant inbreeding depression for seed mass and germination and cumulative early inbreeding depression varied greatly among families. Among sex morphs, we found that females and hermaphrodites differed in biparental inbreeding depression, whereas uniparental was similar for all. Significant inbreeding depression levels may play a role in female maintenance in this species, and individual variation in association with sex-lineages proclivity is discussed.
The genus Silene, studied by Darwin, Mendel and other early scientists, is re-emerging as a system for studying interrelated questions in ecology, evolution and developmental biology. These questions include sex chromosome evolution, epigenetic control of sex expression, genomic conflict and speciation. Its well-studied interactions with the pathogen Microbotryum has made Silene a model for the evolution and dynamics of disease in natural systems, and its interactions with herbivores have increased our understanding of multi-trophic ecological processes and the evolution of invasiveness. Molecular tools are now providing new approaches to many of these classical yet unresolved problems, and new progress is being made through combining phylogenetic, genomic and molecular evolutionary studies with ecological and phenotypic data.
Are parasites always harmful to their hosts? By definition, indeed, but in a few cases and particular environments, hosts experience higher fitness in the presence than in the absence of their parasites. Symbiotic associations form a continuum of interactions, from deleterious to beneficial effects on hosts. In this paper, we investigate the outcome of parasite infection of Arabidopsis thaliana by its natural pathogen Hyaloperonospora arabidopsis. This system exhibits a wide range of parasite impact on host fitness with, surprisingly, deleterious effects on high fecundity hosts and, at the opposite extreme, seemingly beneficial effects on the least fecund one. This phenomenon might result from varying levels of tolerance among host lines and even overcompensation for parasite damage analogous to what can be observed in plant-herbivore systems.
- Oct 2008
Questions: Can trade-offs and genotype-environment interactions maintain variability for fitness-related life-history traits? Hypothesis: Transmission success, the equivalent of fecundity, is traded off against minimizing the latent period, the equivalent of age at maturity. Organisms: the non-lethal parasite Hyaloperonospora Arabidopsis (= H. parasitica) and its host plant Arabidopsis thaliana. Methods: We measured latent period, transmission success and host seed production of all combinations of infections between three parasite strains and three host lines, allowing us to calculate phenotypic correlations between these parasite traits and the relation between host and parasite traits. Conclusions: Infected plants that sporulated more rapidly (short latent period) transmitted their parasites less well, revealing a phenotypic trade-off between these important parasite life-history traits. This phenotypic trade-off may help explain why latent period remains variable in nature and has not achieved a uniformly minimal value.
Cyanogenesis is a widespread chemical defence mechanism in plants against herbivory. However, some specialised herbivores overcome this protection by different behavioural or metabolic mechanisms. In the present study, we investigated the effect of presence or absence of cyanogenic glycosides in birdsfoot trefoil (Lotus corniculatus, Fabaceae) on oviposition behaviour, larval preference, larval development, adult weight and nectar preference of the common blue butterfly (Polyommatus icarus, Lycaenidae). For oviposition behaviour there was a female-specific reaction to cyanogenic glycoside content; i.e. some females preferred to oviposit on cyanogenic over acyanogenic plants, while other females behaved in the opposite way. Freshly hatched larvae did not discriminate between the two plant morphs. Since the two plant morphs differed not only in their content of cyanogenic glycoside, but also in N and water content, we expected these differences to affect larval growth. Contrary to our expectations, larvae feeding on cyanogenic plants showed a faster development and stronger weight gain than larvae feeding on acyanogenic plants. Furthermore, female genotype affected development time, larval and pupal weight of the common blue butterfly. However, most effects detected in the larval phase disappeared for adult weight, indicating compensatory feeding of larvae. Adult butterflies reared on the two cyanogenic glycoside plant morphs did not differ in their nectar preference. But a gender-specific effect was found, where females preferred amino acid-rich nectar while males did not discriminate between the two nectar mimics. The presented results indicate that larvae of the common blue butterfly can metabolise the surplus of N in cyanogenic plants for growth. Additionally, the female-specific behaviour to oviposit preferably on cyanogenic or acyanogenic plant morphs and the female-genotype-specific responses in life history traits indicate the genetic flexibility of this butterfly species and its potential for local adaptation.
- May 2008
Genetic diversity for plant defense against microbial pathogens has been studied either by analyzing sequences of defense genes or by testing phenotypic responses to pathogens under experimental conditions. These two approaches give different but complementary information but, till date, only rare attempts at their integration have been made. Here we discuss the advances made, because of the two approaches, in understanding plant-pathogen coevolution and propose ways of integrating the two.
Bayesian 50% majority-rule consensus tree of the Microbotryum strains analyzed in this study based on the Ef1α gene. Statistical supports indicate Bayesian Posterior Probabilities (BPP)/Maximum Parsimony Bootstraps/Neighbor-Joining Bootstraps. Only nodes supported by more than two methods are indicated, the significant statistical supports being considered as higher than respectively 0.9/70/70. The tree is rooted based on previous studies (see text). Taxa labels correspond to the host plant on which fungal strains were collected. Clades not supported in the individual tree are indicated in grey.
Bayesian 50% majority-rule consensus tree of the Microbotryum strains analyzed in this study based on the γ-tub gene. Statistical supports indicate Bayesian Posterior Probabilities (BPP)/Maximum Parsimony Bootstraps/Neighbor-Joining Bootstraps. Only nodes supported by more than two methods are indicated, the significant statistical supports being considered as higher than respectively 0.9/70/70. The tree is rooted based on previous studies (see text). Taxa labels correspond to the host plant on which fungal strains were collected. Clades not supported in the individual tree are indicated in grey.
Bayesian 50% majority-rule consensus tree of the Microbotryum strains analyzed in this study based on the β-tub gene. Statistical supports indicate Bayesian Posterior Probabilities (BPP)/Maximum Parsimony Bootstraps/Neighbor-Joining Bootstraps. Only nodes supported by more than two methods are indicated, the significant statistical supports being considered as higher than respectively 0.9/70/70. The tree is rooted based on previous studies (see text). Taxa labels correspond to the host plant on which fungal strains were collected. Clades not supported in the individual tree are indicated in grey.
Using phylogenetic approaches, the expectation that parallel cladogenesis should occur between parasites and hosts has been validated in some studies, but most others provided evidence for frequent host shifts. Here we examine the evolutionary history of the association between Microbotryum fungi that cause anther smut disease and their Caryophyllaceous hosts. We investigated the congruence between host and parasite phylogenies, inferred cospeciation events and host shifts, and assessed whether geography or plant ecology could have facilitated the putative host shifts identified. For cophylogeny analyses on microorganisms, parasite strains isolated from different host species are generally considered to represent independent evolutionary lineages, often without checking whether some strains actually belong to the same generalist species. Such an approach may mistake intraspecific nodes for speciation events and thus bias the results of cophylogeny analyses if generalist species are found on closely related hosts. A second aim of this study was therefore to evaluate the impact of species delimitation on the inferences of cospeciation. We inferred a multiple gene phylogeny of anther smut strains from 21 host plants from several geographic origins, complementing a previous study on the delimitation of fungal species and their host specificities. We also inferred a multi-gene phylogeny of their host plants, and the two phylogenies were compared. A significant level of cospeciation was found when each host species was considered to harbour a specific parasite strain, i.e. when generalist parasite species were not recognized as such. This approach overestimated the frequency of cocladogenesis because individual parasite species capable of infecting multiple host species (i.e. generalists) were found on closely related hosts. When generalist parasite species were appropriately delimited and only a single representative of each species was retained, cospeciation events were not more frequent than expected under a random distribution, and many host shifts were inferred.Current geographic distributions of host species seemed to be of little relevance for understanding the putative historical host shifts, because most fungal species had overlapping geographic ranges. We did detect some ecological similarities, including shared pollinators and habitat types, between host species that were diseased by closely related anther smut species. Overall, genetic similarity underlying the host-parasite interactions appeared to have the most important influence on specialization and host-shifts: generalist multi-host parasite species were found on closely related plant species, and related species in the Microbotryum phylogeny were associated with members of the same host clade. We showed here that Microbotryum species have evolved through frequent host shifts to moderately distant hosts, and we show further that accurate delimitation of parasite species is essential for interpreting cophylogeny studies.
- Aug 2007
Congruence between host and parasite phylogenies is often taken as evidence for cospeciation. However, 'pseudocospeciation', resulting from host-switches followed by parasite speciation, may also generate congruent trees. To investigate this process and the conditions favouring its appearance, we here simulated the adaptive radiation of a parasite onto a new range of hosts. A very high congruence between the host tree and the resulting parasite trees was obtained when parasites switched between closely related hosts. Setting a shorter time lag for speciation after switches between distantly related hosts further increased the degree of congruence. The shape of the host tree, however, had a strong impact, as no congruence could be obtained when starting with highly unbalanced host trees. The strong congruences obtained were erroneously interpreted as the result of cospeciations by commonly used phylogenetic software packages despite the fact that all speciations resulted from host-switches in our model. These results highlight the importance of estimating the age of nodes in host and parasite phylogenies when testing for cospeciation and also demonstrate that the results obtained with software packages simulating evolutionary events must be interpreted with caution.
Evolution of parasite traits is inextricably linked to their hosts. For instance one common definition of parasite virulence is the reduction in host fitness due to infection. Thus, traits of infection must be viewed in both protagonists and may be under shared genetic and physiological control. We investigated these questions on the oomycete Hyaloperonospora arabidopsis (= parasitica), a natural pathogen of the Brassicaceae Arabidopsis thaliana. We performed a controlled cross inoculation experiment confronting six lines of the host plant with seven strains of the parasite in order to evaluate genetic variation for phenotypic traits of infection among hosts, parasites, and distinct combinations. Parasite infection intensity and transmission were highly variable among parasite strains and host lines but depended also on the interaction between particular genotypes of the protagonists, and genetic variation for the infection phenotype of parasites from natural populations was found even at a small spatial scale within population. Furthermore, increased parasite fitness led to a significant decrease in host fitness only on a single host line (Gb), although a trade-off between these two traits was expected because host and parasite share the same resource pool for their respective reproduction. We propose that different levels of compatibility dependent on genotype by genotype interactions might lead to different amounts of resources available for host and parasite reproduction. This variation in compatibility could thus mask the expected negative relationship between host and parasite fitness, as the total resource pool would not be constant. These results highlight the importance of host variation in the determination of parasite fitness traits. This kind of interaction may in turn decouple the relationship between parasite transmission and its negative effect on host fitness, altering theoretical predictions of parasite evolution.
- Feb 2007
We investigate the geographic pattern of adaptation of a fungal parasite, Colletotrichum lindemuthianum, on two host species, Phaseolus vulgaris and P. coccineus for two parasite fitness traits: infectivity (ability to attack a host individual) and aggressivity (degree of sporulation and leaf surface damage). Using a cross-inoculation experiment, we show specialization of the fungus on its host species of origin for both traits even when fungi, which originated from hosts growing in sympatry, were tested on sympatric host populations. Within the two host species, we compared infectivity and aggressivity on local versus allopatric plant-fungus combinations. We found evidence for local adaptation for the two traits on P. vulgaris but not on P. coccineus. There was no significant correlation between the degrees of local adaptation for infectivity and aggressivity, indicating that the genetic basis and the effect of selection may differ between these two traits. For the two fitness traits, a positive correlation between the degree of specialization and the degree of local adaptation was found, suggesting that specialization can be reinforced by local adaptation.
- Jun 2006
Microbotryum violaceum is a fungus that causes the sterilizing anther smut disease in Caryophyllaceae. Its diploid teliospores normally produce equal proportions of haploid sporidia of its two mating types. However natural populations contain high frequencies of individuals producing sporidia of only one mating type ('biased strains'). This mating type-ratio bias is caused by deleterious alleles at haploid phase ('haplo-lethals') linked to the mating type locus that can be transmitted only by intra-tetrad selfing. We used experimental inoculations to test some of the hypotheses proposed to explain the maintenance of haplo-lethals. We found a disadvantage of biased strains in infection ability and high intra-tetrad mating rates. Biased strains had no higher competitive ability nor shorter latency and their higher spore production per flower appeared insufficient to compensate their disadvantages. These findings were only consistent with the hypothesis that haplo-lethals are maintained under a metapopulation structure because of high intra-tetrad selfing rates, founder effects and selection at the population level.
Studies of bioinvasions have revealed various strategies of invasion, depending on the ecosystem invaded and the alien species concerned. Here, we consider how migration (as a demographic factor), as well as ecological and evolutionary changes, affect invasion success. We propose three main theoretical scenarios that depend on how these factors generate the match between an invader and its new environment. Our framework highlights the features that are common to, or differ among, observed invasion cases, and clarifies some general trends that have been previously highlighted in bioinvasions. We also suggest some new directions of research, such as the assessment of the time sequence of demographic, genetic and environmental changes, using detailed temporal surveys.
Simultaneous effects of host and parasite in determining quantitative traits of infection have long been neglected in theoretical and experimental investigations of host-parasite coevolution with the notable exception of gene-for-gene resistance studies. A cross-infection experiment, using five lines of the plant Arabidopsis thaliana and two strains of its oomycete pathogen Hyaloperonospora parasitica, revealed that three traits traditionally considered those of the parasite (number of infected leaves, transmission success, and time until 50% transmission), differed among specific combinations of host and parasite lines, being determined by the two protagonists of the infection. However, the two parasite strains did not differ significantly for most measured phenotypic traits of the infection. Globally, transmission increased with increasing virulence among the different host-parasite combinations, as assumed by most models of evolution of virulence. Surprisingly, however, there was no general relationship between parasite and host fitness, estimated respectively as transmission and seed production. Only one of the two strains showed the expected significant negative genetic correlation between these two variables. Our results thus highlight the importance of taking into account both host and parasite genetic variation because their interaction can lead to unexpected evolutionary outcomes.
Ambrosia artemisiifolia is an aggressive North American annual weed, found particularly in sunflower and corn fields. Besides its economic impact on crop yield, it represents a major health problem because of its strongly allergenic pollen. Ragweed was imported inadvertently to Europe in the 18th century and has become invasive in several countries, notably in the Rhône Valley of France. It has recently expanded in both the Provence-Alpes-Côte-d'Azur and Bourgogne regions. As first steps towards understanding the causes and mechanisms of ragweed invasion, genetic variability of French and North American populations was analysed using microsatellites. Overall genetic variability was similar in North America and in the Rhône-Alpes region, but within-population levels of genetic variability were surprisingly lower in native than in invasive French populations. French populations also exhibited lower among-population differentiation. A significant pattern of isolation by distance was detected among North American populations but not among French populations. Assignment tests and distribution of rare alleles did not point to a single origin for all French populations, nor for all individuals within populations and private alleles from different North American populations were found in the same French populations. Indeed, within all French populations, individual plants were roughly equally assigned to the different North American populations. Altogether, these results suggest that the French invasive populations include plants from a mixture of sources. Reduced diversity in populations distant from the original area of introduction indicated that ragweed range expansion probably occurred through sequential bottlenecks from the original populations, and not from subsequent new introductions.
When invading new regions exotic species may escape from some of their natural enemies. Reduced top-down control ("enemy release") following this escape is often invoked to explain demographic expansion of invasive species and also may alter the selective regime for invasive species: reduced damage can allow resources previously allocated to defence to be reallocated to other functions like growth and reproduction. This reallocation may provide invaders with an "evolution of increased competitive ability" over natives that defend themselves against specialist enemies. We tested for enemy release and the evolution of increased competitive ability in the North American native ragweed (Ambrosia artemisiifolia: Asteraceae), which currently is invading France. We found evidence of enemy release in natural field populations from the invaded and native ranges. Further we carried out a reciprocal transplant experiment, comparing several life history traits of plants from two North American (Ontario and South Carolina) and one French population in four common gardens on both continents. French and Canadian plants had similar flowering phenologies, flowering earlier than plants from further south in the native range. This may suggest that invasive French plants originated from similar latitudes to the Canadian population sampled. As with natural populations, experimental plants suffered far less herbivore damage in France than in Ontario. This difference in herbivory translated into increased growth but not into increased size or vigour. Moreover, we found that native genotypes were as damaged as invading ones in all experimental sites, suggesting no evolutionary loss of defence against herbivores.
- Dec 2005
In gynodioecious plant species, females are expected to have more resources available for maturing seeds because pistillate flowers are smaller, do not produce pollen, and are thus less costly that perfect flowers. The potential female advantage arising from more abundant resources is, however, likely to vary depending on whether seed production is limited by resource or pollen availability. Here we experimentally investigated the influence of pollen and resource limitation on female advantage in a gynodioecious species using two levels of pollination. Total seed production of females was always greater than that of hermaphrodites: females produced more flowers and more fruits that contained similar numbers of seeds of similar mass. Under low pollination, female and hermaphrodite plants allocated resources to increased flower production rather than to increased seed size or quality. We did not detect any influence of pollen or resource limitation on female advantage, which remained similar under low (= abundant resources) and full pollination. Outcrossed fruits performed better than selfed fruits when the same plant received both selfed and outcrossed pollen on different flowers. These differences were not greater under high pollination, possibly because resources available for each fruit did not differ between our pollen intensity treatments.
- Oct 2005
The study of how parasites adapt to new hosts is of great importance for understanding the emergence of new diseases. Here, we report a study of the anther smut disease on Gypsophila repens (Caryophyllaceae). In contrast to what is usually found on other host species, infected natural populations of G. repens are extremely rare. Moreover, symptoms of diseased plants are incomplete and highly variable over the time. These results suggest that the fungus infecting G. repens is a case of a parasite not capable of exploiting its host optimally. Molecular analyses of Microbotryum violaceum strains infecting this and other Caryophyllaceae revealed that this sub-optimal behaviour probably resulted from a recent host shift from the morphologically similar plant Petrorhagia saxifraga. With its exceptionally low virulence and prevalence, but apparent self-sustainability, the disease on G. repens may thus represent an interesting case study for investigating the conditions leading to adaptation of parasites on new hosts.
The anther smut fungus Microbotryum violaceum has a mixed mating system, being able to either self or outcross. The infection success may be influenced by selfing or outcrossing. It will affect the amount and distribution of genetic variation within natural populations and, in particular, in this system, may explain the maintenance of common highly deleterious alleles linked to the mating-type locus. Using artificial inoculations with two different mixtures of strains that have different identifiable genotypes on the host plant Silene latifolia, we estimate infection success and selfing propensity of this fungus. Fungal spores were placed in flowers, mimicking natural spore deposition by pollinators. Successful infection was found in 31.5% of inoculated plants, and infection success depended on the number of inoculations and host plant sex, females having a higher per-contact risk but males a higher global probability of infection. Many host individuals were infected by more than one fungal genotype, and separate stems of the same plant were infected independently. We found 14.3% and 28.3% outcrossing events for the two mixtures of fungal strains, confirming that M. violaceum is predominantly selfing. Because we inoculated with spore mixtures, this predominant selfing did not result from a lack of available mating partners, which could further reduce outcrossing in nature. We suggest that selfing may largely result from intratetrad mating (¼intrameiotic), and we estimated the intratetrad mating rate between 43.3% and 71.4%. This has important consequences for understanding the maintenance of haplolethal alleles in M. violaceum.
Females of gynodioecious species need to have reproductive advantages to compensate for their lack of male function and assure their maintenance in natural populations. Females may be more fecund than hermaphrodites because they reallocate resources from male to female function, avoid selfing and inbreeding depression, or produce higher quality offspring by screening arriving pollen better. A novel type of female advantage is proposed that may be important for several species of gynodioecious Caryophyllaceae. The anther smut fungus Microbotryum violaceum generally sterilizes its infected hosts but even without infection, spore deposition can negatively affect the reproduction of healthy individuals. In Gypsophila repens, flowers of hermaphrodite plants experimentally pollinated with both fungal spores and pollen produced significantly fewer fruits and seeds than those that received pollen alone, whereas female reproduction was unaffected by spore deposition. This unexplored reproductive advantage of females is probably due to the larger stigmatic surface in pistillate flowers, that allows pollen germination despite the presence of spores. Because longer stigmas may also lead to increased spore deposition, these results raise questions regarding the infection probability for each sex, possible sex-specific defense mechanisms, and the general role of pathogens in the maintenance of females in natural gynodioecious populations.
tested for sex-differential pollen limitation at the flower level, comparing fruit production and fecundity between open-pollinated and hand-pollinated flowers of Gypsophila repens (Caryophyllaceae) over two years. No sex difference in pollen limitation was found. Hand-pollination increased fruit and seed production in the first year of the study but not in the second, and it actually decreased fruit set in the latter year, although overall fecundity was higher in that year. Ovule number differed between years, with perfect flowers containing more ovules in the second year. Gypsophila repens harbors the anther smut fungus Microbotryum violaceum, which infects and sterilizes many species of Caryophyllaceae. In G. repens, however, some infected plants are capable of seed production, although their fecundity is far lower than that of healthy plants. Seed production was not pollen limited even with the high percentage of infected individuals that could reduce the quantity of pollen available at the population level.
We combined pedigree data with data derived from 14 microsatellite loci to investigate genetic diversity and its maintenance in the captive source population for the reintroduction of the bearded vulture into the Alps. We found the captive population to be genetically more variable than the largest natural population in Europe, both in terms of mean number of alleles per locus and mean observed and expected heterozygosity. Allelic diversity of the captive population was higher than, and mean heterozygosity measurements were comparable with the ones found in two large, extinct populations from Sardinia and the Alps represented by museum specimens. The amount of genetic variability recruited with the founders was still present in the captive population of the year 2000, mainly because the carriers of rare alleles were still alive. However, the decline in expected heterozygosity and the loss of alleles over generations in captivity was significant. Point estimates of effective population size, N(e), based on pedigree data and estimates of effective number of breeders, N(b), based on allele frequency changes, ranged from 20 to 30 and were significantly smaller than the census size. The results demonstrate that the amount of genetic variability in the captive bearded vulture population is comparable or even larger than the amount present in natural populations. However, the population is in danger to lose genetic variability over time because of genetic drift. Management strategies should therefore aim at preserving genetic variability by minimising kinship, and at increasing N(e) by recruiting additional founders and enhancing gene flow between the released, the captive and natural populations.
- Jul 2003
Sexual selection was proposed by Darwin to explain the evolution of traits that do not contribute to survival or fecundity, such as the elaborate train of the peacock, the horn of the rhinoceros beetle, and the bright red belly of the stickleback. The development of such characteristics gives some individuals higher reproductive success because they achieve more matings or successful fertilizations. Sexual selection operates between individuals of the same sex as they compete for mates, and between the sexes as individual choices are made of mating partner.
The present study explores the maintenance of a segregation distortion in the phytopathogenic fungus Microbotryum violaceum (anther smut). Some individuals of this species produce spores with completely biased sex ratio because of the presence of haplo-lethal alleles linked to the mating-type gene. To test the different hypotheses proposed to explain the maintenance of these haplo-lethal alleles (diploid advantage to counter- balance haploid deleterious effects, neutrality of haplo-lethal alleles only in populations with high interstrain competition, and complete neutrality), sex-ratio bias was investigated in natural European populations of M. violaceum from Silene latifolia and other species of Caryophyllaceae. The negative relationship observed between proportions of strains expressing a sex-ratio bias and disease prevalence was opposite the one predicted by the second hypothesis. On the contrary, the pattern of the variances, with proportions of biased strains more extreme in smaller populations, fitted perfectly with the prediction of the third hypothesis. Experimental studies are now needed to confirm the complete neutrality of the haplo-lethal alleles linked to the mating-type gene in M. violaceum.
Polygamy (including trioecy and subdioecy), the co-occurrence of males, hermaphrodites, and females in natural populations, is a rare and poorly studied breeding system expressed in Fraxinus excelsior L. (Oleaceae), a wind-pollinated tree. Here we investigate siring ability of pollen from male vs. hermaphrodite individuals to better understand this sex polymorphism. We conducted single-donor and two-donor pollination experiments and compared both fruit set and seed siring success, assessed with polymorphic microsatellite markers, of male and hermaphrodite individuals. Single pollen donor crosses allowed us to verify the male function of hermaphrodites. However, pollen from hermaphrodites was much less proficient than male pollen, with males siring 10 times as many fruits in single donor pollination treatments. This result was strengthened by the surprisingly low reproductive success of hermaphrodites in pollen competition conditions: of the 110 seedlings analyzed three were selfed and only one was sired by the hermaphrodite donor. The remaining 106 were sired by the male pollen donor. These results raise the question of the maintenance of male fertility in hermaphrodites in Fraxinus excelsior. Male function of hermaphrodites in this species now needs to be assessed under field conditions.
Female fecundity advantage in gynodioecious plants is required for the spread and maintenance of this reproductive system. However, not all reproductive characters show female advantage in all species. We used a meta-analysis to summarise differences between females and hermaphrodites reported from the literature for several reproductive traits. Further we tested three hypotheses, (1) that female plants of species with many ovules produce more seeds per fruit while those with few ovules produce heavier seeds, (2) that females are more pollen limited than hermaphrodites, and (3) that floral sexual size dimorphism is more pronounced in species with few ovules, either because female reproductive success is less limited by pollen availability in such species or because flowers with few ovules require a smaller floral structure to protect the carpels. Overall, females compared to hermaphrodites produced more but smaller flowers, had higher fruit set, higher total seed production, and produced heavier seeds that germinated better. Species with many versus few ovules differed in female advantage for flower size dimorphism, flower number, fruit set and total seed production. However seed size, seed set per fruit and seed germination differences between females and hermaphrodites did not differ significantly between species with few and many ovules. We also found no evidence for differential pollen limitation between females and hermaphrodites. Degree of floral sexual size dimorphism differed significantly between species with few and many ovules. Though pistillate flowers were generally smaller than those of hermaphrodites, species with many ovules showed less difference in flower size between the sexes, suggesting either that the protective role of the perianth constrains the evolution of sexual size dimorphism in species with many ovules or that selection for adequate pollination in species with many ovules impedes the reduction in flower size of females.
Some species described as gynodioecious are truly gynomonoecious-gynodioecious. Three distinct phenotypes may be found in their natural populations-female and hermaphrodite pure-sexed plants bearing either only pistillate or perfect flowers, respectively, and mixed plants bearing both types of flowers. In one such species, Dianthus sylvestris, we investigated mating system parameters using allozyme data. Outcrossing rates and correlations of outcrossed paternity were calculated for the three types of plants and separately for pistillate and perfect flowers on mixed plants. The mean outcrossing rate for the population was t(m) ± SD = 0.885 ± 0.032. Females were more outcrossed than hermaphrodites (0.987 ± 0.112 and 0.790 ± 0.076, respectively), whereas mixed plants were not significantly more or less outcrossed than hermaphrodites (0.840 ± 0.060). Within mixed plants, perfect flowers showed an intermediate outcrossing rate (0.898 ± 0.057), whereas pistillate flowers were as selfed as perfect flowers on hermaphrodite plants (0.782 ± 0.111). Family estimates of outcrossing rates were highly variable. Globally, no biparental inbreeding was detected in this species, and there was a mean of 61.5 ± 19.9% of full-sibs within families. Floral dimorphism between small pistillate and large perfect flowers together with pollinator preference for larger flowers could explain the observed patterns for both mating parameters. The advantages of gynomonoecy-gynodioecy are discussed. We conclude that mixed plants do not reduce selfing for all flowers on a plant, but perfect flowers on these plants seem to have an outcrossing advantage.
- Sep 2002
In spatially structured populations, host–parasite coevolutionary potential depends on the distribution of genetic variation within and among populations. Inoculation experiments using the plant, Silene latifolia, and its fungal pathogen, Microbotryum violaceum, revealed little overall differentiation in infectivity/resistance, latency or spore production among host or pathogen populations. Within populations, fungal strains had similar means, but varied in performance across plant populations. Variation in resistance among seed families indicates the potential for parasite-mediated selection, whereas there was little evidence for local pathogen genotype × plant genotype interactions assumed by most theoretical coevolution models. Lower spore production on sympatric than allopatric hosts confirmed local fungal maladaptation already observed for infectivity. Correlations between infectivity and latency or spore production suggest a common mechanism for variation in these traits. Our results suggest low variation available to this pathogen for tracking its coevolving host. This may be caused by random drift, breeding system or migration characteristic of metapopulation dynamics.
We report the development of 44 microsatellite markers in three host races of the fungus Microbotryum violaceum, a sexually transmitted disease of the Caryophyllaceae. An enrichment protocol was used to isolate microsatellite loci from three strains, collected, respectively, from the plant species Gypsophila repens, Dianthus sylvestris, and Silene vulgaris. Polymorphism and cross-amplification were explored with 32 isolates of M. violaceum, collected on 12 different plant species in natural populations.
Abstract 1. Plant growth and chemical defence compounds in four Lotus corniculatus genotypes exposed to factorial combinations of ambient and elevated carbon dioxide, and herbivory by caterpillars of Polyommatus icarus were measured to test the predictions of the carbon/nutrient balance hypothesis. 2. Shoot and root biomass, allocation to shoots versus roots, and carbon-based defence compounds were greater under elevated carbon dioxide. Pupal weight of P. icarus was greater and development time shorter under elevated carbon dioxide. 3. Herbivory decreased shoot growth relative to root growth and production of nitrogen-based defence (cyanide). Young leaves contained more defence compounds than old leaves, and this response depended on carbon dioxide and herbivory treatments (significant interactions). 4. Genotype-specific responses of plants to carbon dioxide and herbivory were found for the production of cyanide. Furthermore, maternal butterfly-specific responses of caterpillars to carbon dioxide were found for development time. This suggests the existence of genetic variation for important defence and life-history traits in plants and herbivores in response to rising carbon dioxide levels.
- Apr 2002
1. Four Lotus corniculatus genotypes differing in cyanoglycoside and condensed tannin concentrations were grown in either low (350 ppm) or high (700 ppm) atmospheric CO2 environments. Larval performance, consumption and conversion efficiency of Polyommatus icarus feeding on this plant material were measured. 2. Plants grown under elevated CO2 contained less cyanoglycosides, more condensed tannins and more starch than control plants. However, water concentration, nitrogen and protein as well as nitrogen concentration in relation to carbon concentration did not differ between CO2 treatments. 3. The four genotypes differed significantly in condensed tannins, cyanoglucoside, leaf water and leaf nitrogen but no genotype–CO2 interaction was detected, except for total phenolics and condensed tannins in which two plant genotypes showed stronger increases under elevated CO2 than the other two. 4. Larvae of P. icarus consumed more plant material and used and converted it more efficiently from plants grown at high atmospheric CO2. 5. Larvae developed significantly faster and were significantly heavier when fed plant material grown under elevated CO2. The observed difference in mass disappeared in the pupal and adult stages. However, lipid concentration of adults from the elevated CO2 treatment was marginally significantly higher than of controls. 6. It is concluded that the higher carbohydrate concentration of L. corniculatus plants grown at elevated CO2 renders leaves more suitable and better digestible to P. icarus. Furthermore, differences in allelochemicals might influence the palatability of L. corniculatus leaves for this specialist on Fabaceae.
In insect-pollinated plants flowers must balance the benefits of attracting pollinators with the cost of attracting natural enemies, when these respond to floral traits. This dilemma can have important evolutionary consequences for mating-system evolution and polymorphisms for floral traits. We investigate the benefits and risks associated with flower size and sex morph variation in Dianthus sylvestris, a gynodioecious species with pistillate flowers that are much smaller than perfect flowers. We found that this species is mainly pollinated by nocturnal pollinators, probably moths of the genus Hadena, that also oviposit in flowers and whose caterpillars feed on developing fruits and seeds. Hadena preferred larger flowers as oviposition sites, and flowers in which Hadena had deposited eggs bore more pollen on their stigmas, suggesting that Hadena is indeed the principle pollinator, or that pollinators and these seed predators employ the same choice criteria for flowers. Globally, perfect flowers suffered more predation by seed predators than did pistillate flowers, suggesting that seed predators play an important role in reproductive system dynamics in this species. On the other hand, female flowers were more likely to be contaminated with spores of another natural enemy, the pathogenic fungus Microbotryum violaceum, that are transmitted by pollinating insects. This complex interplay between two natural enemies, one of which, Hadena, is also a pollinator and vector of fungal spores, may contribute to the maintenance of the polymorphic reproductive system of the host plant.
- Dec 2001
Summary • Behavioural, physiological or immunological constraints often render one sex more susceptible to parasites, thereby potentially generating sex-specific trade-offs between traits associated with infection risk and other life-history characters. • The fungal pathogen Microbotryum violaceum systemically infects the dioecious plant Silene latifolia when pollinators deposit fungal spores on the flowers of healthy plants. Male plants produce many short-lived flowers, whereas females produce few flowers that remain connected with the plant after fertilization. We investigated how variation in flower production and flower longevity affects the infection risk for males and females. • In glasshouse experiments, we varied the number of flowers inoculated (4 vs. 16 per plant) with spores and the time until these flowers were removed (1 or 2 days for both sexes, 14 days for females only). We also measured the longevity of male flowers receiving simulated visits, with or without spores, to test for an abscission response to visitation and/or contamination. In a field survey, we measured male and female disease prevalence in 17 natural populations. • Varying the number of inoculated flowers did not affect infection probability, but females retaining inoculated flowers for 14 days became diseased more often (20.0%) than did plants with flowers removed within 2 days (7.3%). • Males that had dropped more inoculated flowers prematurely were more likely to remain uninfected. Spore-bearing visits shortened male flower longevity (38.4 ± 2.8 h) relative to non-spore visits (47.9 ± 5.2 h). • Female field disease prevalence (19.7 ± 3.5%) was higher than that of males (14.3 ± 2.6%), especially in populations with a high disease incidence. • Continuing physical connection during fruit ripening appears to increase invasion time and thus the per-contact infection risk in females. This is consistent with higher female field prevalences, although other explanations, unrelated to disease transmission, are possible. These results illustrate how interactions between plant reproductive behaviour and pollinator activity may affect disease spread. Female mating behaviour may evolve towards lower attractiveness to pollinators to minimize infectious contacts, while males can afford to be more promiscuous with an attractive, but disposable, floral display.
- Mar 2001
We investigated the genetic population structure of the sexually transmitted plant pathogen, the fungus Microbotryum violaceum, on the two closely related host species Silene latifolia and S. dioica using microsatellite markers. We found strong deviations from Hardy-Weinberg expectations, with significant heterozygote deficiency in almost all populations. Fungal strains from the two host species were differentiated, and these host races differed in amount of variation within populations and differentiation among populations. Anther smut from S. latifolia harboured significantly less microsatellite diversity and were more genetically differentiated from each other than those from S. dioica. Small effective population sizes, rapid population turnover, and less gene flow among populations could lead to this higher population differentiation and lower within population genetic diversity for anther smut populations on S. latifolia than on S. dioica. These results are in concordance with host ecology because S. latifolia grows in more disturbed habitats than S. dioica and may provide a shorter-lived host environment.
Fluctuating asymmetry (FA) is considered to be a good measure of developmental stability. We measured the asymmetry of leaves and flowers of 16 different genotypes of Lotus corniculatus grown in four different experimental environments to estimate the plasticity or developmental stability of asymmetry itself. We found that an index of FA (absolute difference between size of left and right sides, corrected for trait size) differed significantly across environments, with the treatment CO2+/N+ inducing the greatest FA for both flowers and leaves. Genotypes did not differ in FAs. Individual plants showed significantly different FAs only for flowers. At the individual level, we found no significant relationship between flower FA and fitness. Previous work indicates that change in asymmetry in a poor or perturbing environment versus a good environment could reflect the intrinsic quality of a particular genotype. However, in our experiment, genotype effect was significant only for change in asymmetry of leaves, and this last trait was not significantly correlated with our fitness estimate for each genotype in either the most or the least perturbing environment.
- Mar 2000
The anther smut fungus Microbotryum violaceum ( = Ustilago violacea) is a parasite of many species of the Caryophyllaceae. Its host specificity has been debated since early this century, when cross-inoculation experiments indicated the existence of host-specific lineages. Recently, on the basis of spore ultrastructure, all presumed host races were lumped within M. violaceum. To measure gene flow among natural populations of anther smuts from different host species, we used microsatellite variation at 5 loci among samples from 8 Silene, 2 Saponaria, 2 Dianthus and 1 Gypsophila species. Most of the 326 M. violaceum samples investigated originated from the Swiss Alps and close surroundings. Microsatellite variation revealed almost perfect isolation among anther smut fungi from different host species. In addition, differentiation was supported by the nonrandom distribution of null alleles among samples from different host species and host genera. Null alleles were most abundant in anther smut samples from non-Silene hosts. The resolution of genetic differentiation among anther smuts from different host species was highest in those from Silene species. Genetic relationships among samples as indicated by Neighbour-Joining analysis based on genetic distances are discussed with respect to host phylogeny and host ecology. One sample was identified as Ustilago gausseni because it had verrucose instead of reticulate spores and was collected from Silene italica. Neighbour-Joining analysis revealed that this sample was similar to the M. violaceum samples from other Silene host species. Therefore, our data question spore morphology as a reliable character for anther smut systematics.
- Dec 1999
Minor bilateral or radial asymmetry of leaves or flowers, the frequency of phenodeviants, intraindividual variation in repeated characters, and fractal dimensions of morphology are considered to represent measures of developmental instability since deviations from regularity of the phenotype constitute a measure of the inability to maintain developmental precision during ontogeny. First, we review patterns of fluctuating asymmetry in plants and show that levels of asymmetry are considerably greater than in animals. While petal asymmetry tends to decrease with petal size within species, leaf asymmetry tends to increase with leaf size. Intraspecific correlations of petal asymmetry and leaf asymmetry are weakly positive. Second, a meta-analysis of the effects of environmental factors hypothesized to increase asymmetry in leaves and flowers, such as radiation, ultraviolet light, excess artificial fertilizer, pollutants, extreme saline conditions, herbivory, and competition, showed intermediate (i.e., explaining 10% of the variance) to large (i.e., explaining 25% of the variance) effects. Third, a meta-analysis of the effects of genetic factors hypothesized to contribute to increased asymmetry in plants, such as homozygosity, hybridization, mutation, and quantitative genetic differences among individuals, showed variable but usually significant effects, although the number of studies generally was small. Controlled experimental studies of environmental and genetic effects on developmental instability of plants may increase our understanding of the mechanisms causing developmental instability.
Genetic isolation among strains of the plant pathogenic fungus Microbotryum violaceum on three species of its host plants was examined. Fungal strains collected from a sympatric population of all three host plant species were examined for their ability to infect the other host plant species and to cross-conjugate among each other. Genetic isolation was investigated from the distribution of neutral microsatellite alleles. Since this is a pollinator-transmitted disease, we examined movement patterns and flower-visitation behavior of pollinators to investigate whether they transfer fungal spores between different host species. Low infection success from the cross-inoculation experiment limits interpretability of the results, but fungus collected from Silene vulgaris was capable of infecting Dianthus carthusianorum. Different fungal strains were able to conjugate and form the infectious dikaryon in most combinations, so hybridization between different fungal host races is possible. The distribution of neutral genetic variation, however, revealed little successful genetic exchange among the fungal host races that were clearly differentiated by host plant species. Pollinators, while showing partial constancy, moved between plants of different host species. Pollinator behavior is therefore not adequate to explain the lack of gene flow among the different fungal races. This indicates that the divergence among these fungal races that has produced incipient species sharing almost no alleles may have occurred in allopatry, unless disruptive selection can outweigh gene flow among fungal races in sympatry.
Evidence is accumulating about the way strategic decisions over an organism's lifetime may depend on its state and on external circumstances. In the parasitic copepod Pachypygus gibber, sex is environmentally determined when the free-living infective stage (Copepodid 2) enters the host, the sea squirt Ciona intestinalis. Whereas in a rich trophic environment the Copepodid 2 develops into a typical male or into a typical female (both are unable to swim and must remain inside the host), in poor trophic conditions it develops into an alternative free-living stage able to swim and to leave the host, the atypical male. Thus this copepod life cycle is characterised by 2 fundamentally different free-living stages. We conducted an experimental study to compare the priority decisions of the Copepodid 2 and of the atypical male in their habitat selection behaviour. We showed that while Copepodid 2s are mainly attracted by chemicals released by the host into the water, atypical males only respond to female and typical male cues. We discuss the adaptiveness on this behavioural difference in light of the ecological constraints met by the different stages of P. gibber.
- Jul 1999
Individual differences in developmental stability are often estimated using individual asymmetry for bilaterally symmetric traits. These are expected to reflect intrinsic abilities to cope with adverse environmental conditions, an important aspect of genotypic quality which should be associated with fitness, but this supposition has never been tested. Two experiments which manipulated environmental conditions were used to investigate whether individual asymmetry under good environmental conditions adequately predicts a genotype's capacity to buffer development across environments, and further to test whether individuals with superior buffering capacity against environmental stress, as reflected in less increase in asymmetry between poor and good environments, had higher fitness than those with poor buffering capacity, after controlling for the influence of asymmetry per se on fitness. Individual asymmetry of the outer tail feathers of male barn swallows, Hirundo rustica, was compared between two different kinds of good and poor environments. Although some individuals changed ranks for asymmetry values between environmental qualities, individual asymmetry under good conditions correlated with this trait in poor environmental conditions. Furthermore, asymmetry under good environmental conditions was positively correlated to change in asymmetry in one of the two experiments, indicating that individual asymmetry is a good predictor of developmental buffering ability. Individuals that increased in asymmetry less under poor environmental conditions had higher reproductive success, even after controlling for asymmetry phenotype, indicating that developmental stability reflects individual quality, as expressed in high relative fitness. These experiments demonstrate that intrinsic differences in developmental stability reliably predict an important fitness component.
We investigated the genetic population structure in a metapopulation of the plant Silene latifolia (Caryophyllaceae) and its fungal pathogen Microbotryum violaceum (Ustilaginales), a pollinator-borne disease. Population structure of the host plant was estimated using allozyme markers and that of the fungus by microsatellites. Both host and parasite showed significant differentiation, but parasite populations were 12 times more strongly differentiated than those of the hosts. We found significant isolation by distance for host populations but not for parasite populations. Higher population differentiation for the parasite may result from small effective population size, high selfing rates, or low migration rate. In this system, hosts are obligate outcrossers and they migrate by seeds and pollen, whereas parasites can self-fertilize and migrate only on pollinating insects. We discuss the effect of limited gene flow in this parasite on its coevolutionary interaction with its host, and its potential for local adaptation on sympatric host populations.
- May 1999
In Thymus vulgaris L., sex determination involves both the nuclear and the cytoplasmic genomes: the cytoplasm is responsible for male-sterility (the female phenotype) while specific nuclear genes may restore male fertility (the hermaphrodite phenotype). Previous observations have shown high variation among hermaphrodites for pollen and seed production. In order to investigate the origin of this variation, 12 female plants, four from each of three populations, were hand-pollinated with pollen from hermaphrodites from three different paternal populations. The sex-ratio (i.e. the frequency of hermaphrodites) produced and the reproductive functions of these offspring were measured. A strong positive correlation was observed between the sex-ratio within a family and both female and male reproductive functions of its hermaphrodites. No such correlation was found for females. This result suggests that restorer genes may be directly or indirectly involved both in sex determination and in the efficiency of resource allocation to reproductive functions. As a consequence, female advantage, i.e. the relative fecundity of females to hermaphrodites, is larger in families with low sex-ratio, and this might affect the evolution of this gynodioecious breeding system.
Habitat fragmentation on the island of of La Réunion (Mascarene Archipelago, Indian Ocean) has increased over three centuries and has now reached a critical level within forest formations at low altitude. This study was performed to examine the consequences of this fragmentation on the threatened species Dombeya acutangula ssp. acutangula. Seven sites of varying population sizes and levels of disturbance were studied. Richness and abundance of all woody species were quantified and reproductive traits of D. acutangula individuals were measured. The smaller proportion of juveniles observed in small populations of D. acutangula might result from fragmentation of natural populations and/or invasion by introduced species. This study moreover shows that D. acutangula exhibited a highly significant decrease in seed production per fruit with decreasing population size, and that pollination was more limited in small populations. We found no evidence of differences among populations for most other traits, particularly on progenies grown under greenhouse conditions. Results are interpreted as a consequence of different, but not mutually exclusive processes, particularly competition from other species and a decrease in availability of compatible pollen in small populations. We suggest several management measures to slow the decline of this species in its natural habitat. These include control measures for invading species and reinforcement of particularly threatened populations through hand-pollination and introduction of individuals with appropriate alleles.
The anther smut fungus Microbotryum violaceum is a pollinator-transmitted plant disease. As for other vector-borne diseases, frequency-dependent transmission patterns are predicted, in contrast to the density-dependent transmission of passively spread diseases. Frequency dependence will, however, only arise if vectors compensate for varying plant spacings. To test this assumption, we set up experimental populations of the host plant, Silene latifolia, with varying disease density (number of diseased plants per plot) and frequency (proportion of plants diseased), and three different plant spacings. We measured spore deposition on healthy flowers in these plots on two dates. Spore deposition decreased considerably from the first to the second census, perhaps related to the concomitant decrease in inflorescence sizes of diseased plants. At our first census, spore deposition rates varied with disease frequency, and the effect of frequency depended on plant spacing. While spore deposition was positively frequency dependent at the 1.5-m inter-plant spacing, no effect of disease frequency was found at a spacings of 0.5 m or 3 m. Nor was there an effect of disease density on spore deposition at the first census. At the later census, on the other hand, spore deposition increased almost significantly with increasing disease density (P = 0.08). This difference in deposition pattern together with a significant decrease in spore receipt indicates changes in pollinator spectrum and/or activity. The correlation of spore numbers among flowers within plants, an indication for intra-plant moves by vectors, was significant at 0.5 m and 1.5 m but not at 3 m. Floral traits and sex of individual plants influenced the number of spores they received. On the first census date, spore deposition increased with increasing inflorescence size in female but not in male plants. On the second census date, neither sex nor number of open flowers had an effect on spore receipt. None of the experimental plants became infected, however, probably because of the unusually hot and dry weather.
Conventional wisdom holds that parasites evolve more rapidly than their hosts and are therefore locally adapted, that is, better at exploiting sympatric than allopatric hosts. We studied local adaptation in the insect-transmitted fungal pathogen Microbotryum violaceum and its host plant Silene latifolia. Infection success was tested in sympatric (local) and allopatric (foreign) combinations of pathogen and host from 14 natural populations from a metapopulation. Seedlings from up to 10 seed families from each population were exposed to sporidial suspensions from each of four fungal strains derived from the same population, from a near-by population (
Conventional wisdom holds that parasites evolve more rapidly than their hosts and are therefore locally adapted, that is, better at exploiting sympatric than allopatric hosts. We studied local adaptation in the insect-transmitted fungal pathogen Microbotryum violaceum and its host plant Silene latifolia. Infection success was tested in sympatric (local) and allopatric (foreign) combinations of pathogen and host from 14 natural populations from a metapopulation. Seedlings from up to 10 seed families from each population were exposed to sporidial suspensions from each of four fungal strains derived from the same population, from a near-by population (< 10 km distance), and from two populations at an intermediate (< 30 km) and remote (< 170 km) distance, respectively. We obtained significant pathogen X plant interactions in infection success (proportion of diseased plants) at both fungal population and strain level. There was an overall pattern of local maladaptation of this pathogen: average fungal infection success was significantly lower on sympatric hosts (mean proportion of diseased plants = 0.32 ± 0.03 SE) than on allopatric hosts (0.40 ± 0.02). Five of the 14 fungal populations showed no strong reduction in infection success on sympatric hosts, and three even tended to perform better on sympatric hosts. This pattern is consistent with models of time-lagged cycles predicting patterns of local adaptation in host-parasite systems to emerge only on average. Several factors may restrict the evolutionary potential of this pathogen relative to that of its host. First, a predominantly selfing breeding system may limit its ability to generate new virulence types by sexual recombination, whereas the obligately outcrossing host 5. latifolia may profit from rearrangement of resistance alleles by random mating. Second, populations often harbor only a few infected individuals, so virulence variation may be further reduced by drift. Third, migration rates among host plant populations are much higher than among pathogen populations, possibly because pollinators prefer healthy over diseased plants. Migration among partly isolated populations may therefore introduce novel host plant resistance variants more often than novel parasite virulence variants. That migration contributes to the coevolutionary dynamics in this system is supported by the geographic pattern of infectivity. Infection success increased over the first 10-km range of host-pathogen population distances, which is likely the natural range of gene exchange.
- Mar 1999
In the fungal pathogen Microbotryumviolaceum mating (i.e. conjugation between cells of opposite mating type) is indispensable for infection of its host plant Silenelatifolia. Since outcrossing opportunities are potentially rare, selfing may be appropriate to ensure reproduction. On the other hand, outcrossing may create genetic variability necessary in the coevolutionary arms race with its host. We investigated the propensity of M. violaceum to outcross vs. self in different host environments. We used haploid sporidia from each of three strains from five fungal populations for pairwise mixtures of opposite mating type, representing either selfing or outcrossing combinations. Mixtures were exposed to leaf extract from seven S. latifolia plants. The proportion of conjugated sporidia quantified mating propensity. The identity of both fungal strains and host influenced conjugation. First, individual strains differed in conjugation frequency by up to 30%, and strains differed in their performance across the different hosts. Second, selfing combinations produced, on average, more conjugations than did outcrossing combinations. Selfing appears to be the predominant mode of reproduction in this fungus, and selfing preference may have evolved as a mechanism of reproductive assurance. Third, individual strains varied considerably in conjugation frequency in selfing and outcrossing combinations across different hosts. This indicates that conjugation between outcrossing partners could be favoured at least in some hosts. Since the dikaryon resulting from conjugation is the infectious unit, conjugation frequency may correspond with infection probability. This assumption was supported by an inoculation experiment, where high infectious sporidial dosage resulted in higher infections success than did low dosage. We therefore predict that sexual recombination can provide this pathogen with novel genotypes able to infect local resistant hosts.
All co-authors (50)