Two colonisation stages generate two different patterns of genetic diversity within native and invasive ranges of Ulex europaeus.

Genetic diversity and the way a species is introduced influence the capacity of populations of invasive species to persist in, and adapt to, their new environment. The diversity of introduced populations affects their evolutionary potential, which is particularly important for species that have invaded a wide range of habitats and climates, such as European gorse, Ulex europaeus. This species originated in the Iberian peninsula and colonised Europe in the Neolithic; over the course of the past two centuries it was introduced to, and has become invasive in, other continents. We characterised neutral genetic diversity and its structure in the native range and in invaded regions. By coupling these results with historical data, we have identified the way in which gorse populations were introduced and the consequences of introduction history on genetic diversity. Our study is based on the genotyping of individuals from 18 populations at six microsatellite loci. As U. europaeus is an allohexaploid species, we used recently developed tools that take into account genotypic ambiguity. Our results show that genetic diversity in gorse is very high and mainly contained within populations. We confirm that colonisation occurred in two stages. During the first stage, gorse spread out naturally from Spain towards northern Europe, losing some genetic diversity. During the second stage, gorse was introduced by humans into different regions of the world, from northern Europe. These introductions resulted in the loss of rare alleles but did not significantly reduce genetic diversity and thus the evolutionary potential of this invasive species.

Genetic variation in flowering phenology and avoidance of seed predation in native populations of Ulex europaeus.

The genetic variation in flowering phenology may be an important component of a species' capacity to colonize new environments. In native populations of the invasive species Ulex europaeus, flowering phenology has been shown to be bimodal and related to seed predation. The aim of the present study was to determine if this bimodality has a genetic basis, and to investigate whether the polymorphism in flowering phenology is genetically linked to seed predation, pod production and growth patterns. We set up an experiment raising maternal families in a common garden. Based on mixed analyses of variance and correlations among maternal family means, we found genetic differences between the two main flowering types and confirmed that they reduced seed predation in two different ways: escape in time or predator satiation. We suggest that this polymorphism in strategy may facilitate maintain high genetic diversity for flowering phenology and related life-history traits in native populations of this species, hence providing high evolutionary potential for these traits in invaded areas.

A model simulating the dynamics of plant mitochondrial genomes.

Molecular evolution of the plant mitochondrial genome involves rearrangements due to the presence of highly recombining repeated sequences. As a result, this genome is composed of a set of molecules of various sizes that generate each other through recombination. The model presented simulates the evolution of various frequencies of the different types of molecules over successive cell cycles. It considers the mitochondrial genome as a population of circular molecules evolving through recombination, replication and random segregation. The model parameters are the rates of recombination of each sequence, the frequency of each type of recombination, the replication rates of the circles and the total amount of mitochondrial DNA per cell. This model demonstrates that high recombination rates lead to rapid deletions of sequences in the absence of selection. The frequency of deletion is dependent on the simulated reproductive mechanism. The conditions leading to reversible or irreversible rearrangements were also investigated.

Variability within the Seychelles cytoplasmic incompatibility system in Drosophila simulans.

In Drosophila simulans, we described a cytoplasmic incompatibility (CI) system (Seychelles) restricted to insular populations that harbor the mitochondrial type SiI. Since then, these populations have been shown to be heterogeneous, some being infected by one Wolbachia genetic variant only (wHa), while others are infected simultaneously by wHa and by another variant (wNo) always found in association with wHa. We have experimentally obtained two D. simulans strains only infected by the wNo variant. This variant determines its own cytoplasmic incompatibility type. In particular, the cross between wNo-bearing flies and wHa-bearing ones is bidirectionally incompatible. The Seychelles CI type, stricto sensu, is distinguished by being determined by the simultaneous presence of two Wolbachia variants that we found to be mutually incompatible. In addition, we observed incomplete maternal transmission of the Wolbachia.

Dynamics of plant mitochondrial genome: model of a three-level selection process.

The plant mitochondrial genome is composed of a set of molecules of various sizes that generate each other through recombination between repeated sequences. Molecular observations indicate that these different molecules are present in an equilibrium state. Different compositions of molecules have been observed within species. Recombination could produce deleted molecules with a high replication rate but bearing little useful information for the cell (such as "petite" mutants in yeast). In this paper we use a multilevel model to examine selection among rapidly replicating incomplete molecules and relatively slowly replicating complete molecules. Our model simulates the evolution of mitochondrial information through a three-level selection process including intermolecular, intermitochondrial, and intercellular selection. The model demonstrates that maintenance of the mitochondrial genome can result from multilevel selection, but maintenance is difficult to explain without the existence of selection at the intermitochondrial level. This study shows that compartmentation into mitochondria is useful for maintenance of the mitochondrial information. Our examination of evolutionary equilibria shows that different equilibria (with different combinations of molecules) can be obtained when recombination rates are lower than a threshold value. This may be interpreted as a drift-mutation balance.

The Y chromosomes of Drosophila simulans are highly polymorphic for their ability to suppress sex-ratio drive.

The sex-ratio trait, known in several species of Drosophila including D. simulans, results from meiotic drive of the X chromosome against the Y. Males that carry a sex-ratio X chromosome produce strongly female-biased progeny. In D. simulans, drive suppressors have evolved on the Y chromosome and on the autosomes. Both the frequency of sex-ratio X and the strength of the total drive suppression (Y-linked and autosomal) vary widely among geographic populations of this worldwide species. We have investigated the pattern of Y-linked drive suppression in six natural populations representative of this variability. Y-linked suppressors were found to be a regular component of the suppression, with large differences between populations in the mean level of suppression. These variations did not correspond to differences in frequency of discrete types of Y chromosomes, but to a more or less wide continuum of phenotypes, from nonsuppressor to partial or total suppressor. We concluded that a large diversity of Y-linked suppressor alleles exists in D. simulans and that some populations are highly polymorphic. Our results support the hypothesis that a Y-chromosome polymorphism can be easily maintained by a balance between meiotic drive and the cost of drive suppression.

Sex-ratio distorter of Drosophila simulans reduces male productivity and sperm competition ability.

The aim of the present study was to determine whether the effects of sex-ratio segregation distorters on the fertility of male Drosophila simulans can explain the contrasting success of these X-linked meiotic drivers in different populations of the species. We compared the fertility of sex-ratio and wild-type males under different mating conditions. Both types were found to be equally fertile when mating was allowed, with two females per male, during the whole period of egg laying. By contrast sex-ratio males suffered a strong fertility disadvantage when they were offered multiple mates for a limited time, or in sperm competition conditions. In the latter case only, the toll on male fertility exceeded the segregation advantage of the distorters. These results indicate that sex-ratio distorters can either spread or disappear from populations, depending on the mating rate. Population density is therefore expected to play a major role in the evolution of sex-ratio distorters in this Drosophila species.

Genetic conflicts.

Self-promoting elements (also called ultraselfish genes, selfish genes, or selfish genetic elements) are vertically transmitted genetic entities that manipulate their "host" so as to promote their own spread, usually at a cost to other genes within the genome. Examples of such elements include meiotic drive genes and cytoplasmic sex ratio distorters. The spread of a self-promoting element creates the context for the spread of a suppressor acting within the same genome. We may thus say that a genetic conflict exists between different components of the same genome. Here we investigate the properties of such conflicts. First we consider the potential diversity of genomic conflicts and show that every genetic system has potential conflicts. This is followed by analysis of the logic of conflicts. Just as Evolutionarily Stable Strategy (ESS) terminology provides a short cut for discussion of much in behavioral ecology, so the language of modifier analysis provides a useful terminology on which to base discussions of conflicts. After defining genetic conflict, we provide a general analysis of the conflicting parties, and note a distinction between competing and conflicting genes. We then provide a taxonomy of possible short- and long-term outcomes of conflicts, noting that potential conflict in an unconstrained system can never be removed, and that the course of evolution owing to conflict is often unpredictable. The latter is most particularly true for strong conflicts in which suppressors may take surprising forms. The possibility of extended conflicts in the form of "arms races" between element and suppressor is illustrated. The peculiar redundancy of these systems is one possible trace of conflict, and others are discussed. That homologous conflicts may find highly different expression is discussed by referring to the mechanistic differences that are thought to underlie the action of the two best-described meiotic drive genes, and by the multiplicity of forms of cytoplasmic sex ratio distorters. The theoretical analysis establishes a logical basis for thinking about conflicts, but fails to establish the importance of conflict in evolution. We illustrate this contentious issue through consideration of some phenomena for whose evolution conflict has been proposed as an important force: the evolution of sex, sex determination, species, recombination, and uniparental inheritance of cytoplasmic genes. In general, it is proposed that conflict may be a central force in the evolution of genetic systems. We conclude that an analysis of conflict and its general importance in evolution is greatly aided by application of the concept of genetic power. We consider the possible components of genetic power and ask whether and how power evolves.

Capillary zone electrophoresis in normal or reverse polarity separation modes for the analysis of hydroxy acid oligomers in neutral phosphate buffer.

Capillary zone electrophoresis (CZE) with neutral phosphate buffer as the background electrolyte was used to analyse water-soluble oligomers obtained by polycondensation of racemic lactic acid. Two CZE separation modes were tested. The first mode was based on normal separation (injection at the anodic side) using a fused-silica capillary. Eight peaks were observed within a 60-min migration time range. They were ascribed to dimer and higher water-soluble oligomers. Peaks from dimer to tetramer were split due to sensitivity for the fine structures at the level of the distribution of chiral lactic acid moieties in oligomer chains. The second mode was based on reverse separation (injection at the cathodic side) using a fused-silica capillary modified by adsorption of a polycation on its inner wall. Under these conditions, oligomers were rapidly separated without peak splitting. Considering the forces which are involved in CZE, data were plotted as a function of 1/t scale, according to the equation [signal]=f((-1)(k)/t) where k=0 and k=1 for normal and reverse separation modes, respectively. Such a plot allowed direct comparison between the various runs after a simple translation along the 1/t axis, regardless of the separation mode and the variation of electroosmotic flow. The second separation mode allowed separation of 3-hydroxybutyric acid and 6-hydroxyhexanoic acid oligomers. For the former series of oligomers, a side reaction generating crotyl bonds was observed due to the high sensitivity of CZE. It was shown that separation was governed by the ratio charge/mass of the oligoesters whatever their structure.

Complex determination of male sterility in Thymus vulgaris L.: genetic and molecular analysis.

Nucleocytoplasmic determination of male sterility in Thymus vulgaris L. has been assumed in all papers attempting to explain the remarkably high frequencies of male steriles found in natural populations of this species. This paper provides strong evidence that both nuclear and cytoplasmic genes are involved in the determination of male sterility of this species, giving a complex inheritance. Interpopulation and intrapopulation crosses have shown that the ratio of females versus hermaphrodites among offsprings varied widely from 1∶0 to 1∶1. Furthermore, interpopulation crosses consistently yielded a higher frequency of females than intrapopulation crosses. Nucleocytoplasmic inheritance was demonstrated by an absence of male fertiles in backcrosses and asymmetrical segregation in reciprocal crosses. Molecular analysis of the mitochondrial DNA of some of the parents used in crosses suggested the involvement of different cytoplasms in the inheritance of male steriliy.