Evidence for the circadian gene period as a proximate mechanism of protandry in a pollinating fig wasp

Protandry in insects is the tendency for adult males to emerge before females and usually results from intra-sexual selection. However, the genetic basis of this common phenomenon is poorly understood. Pollinating fig wasp (Agaonidae) larvae develop in galled flowers within the enclosed inflorescences (‘figs’) of fig trees. Upon emergence, males locate and mate with the still galled females. After mating, males release females from their galls to enable dispersal. Females cannot exit galls or disperse from a fig without male assistance. We sampled male and female Ceratosolen solmsi (the pollinator of Ficus hispida) every 3 h over a 24 h emergence period, and then measured the expression of five circadian genes: period (per), clock (clk), cycle (cyc), pigment-dispersing factor (pdf) and clockwork orange (cwo). We found significant male-biased sexual dimorphism in the expression of all five genes. per showed the greatest divergence between the sexes and was the only gene rhythmically expressed. Expression of per correlated closely with emergence rates at specific time intervals in both male and female wasps. We suggest that this rhythmical expression of per may be a proximate mechanism of protandry in this species.     

Symbiosis as a source of selectable epigenetic variation: taking the heat for the big guy

Evolutionary developmental biology is based on the principle that evolution arises from hereditable changes in development. Most of this new work has centred on changes in the regulatory components of the genome. However, recent studies (many of them documented in this volume) have shown that development also includes interactions between the organism and its environment. One area of interest concerns the importance of symbionts for the production of the normal range of phenotypes. Many, if not most, organisms have ‘outsourced’ some of their developmental signals to a set of symbionts that are expected to be acquired during development. Such intimate interactions between species are referred to as codevelopment, the production of a new individual through the coordinated interactions of several genotypically different species. Within the past 2 years, several research programmes have demonstrated that such codevelopmental schemes can be selected. We will focus on symbioses in coral reef cnidarians symbiosis, pea aphids and cactuses, wherein the symbiotic system provides thermotolerance for the composite organism.     

Rapid evolution of outer egg membrane proteins in the Drosophila melanogaster subgroup: a case of ecologically driven evolution of female reproductive traits

Although sexual selection has been predominantly used to explain the rapid evolution of sexual traits, eggs of oviparous organisms directly face both the challenges of sexual selection as well as natural selection (environmental challenges, survival in niches, etc.). Being the outermost membrane in most insect eggs, the chorion layer is the interface between the embryo and the environment, thereby serving to protect the egg. Adaptive ecological radiations such as divergence in ovipositional substrate usage and host-plant specializations can therefore influence the evolution of eggshell proteins. We can hypothesize that proteins localized on the outer eggshell may be affected to a greater degree by ecological challenges compared with inner eggshell proteins, and therefore, proteins localized in the outer eggshell (chorion membrane) may evolve differently (faster) than proteins localized in the inner egg membrane (vitelline membrane). We compared the evolutionary divergence of vitelline with chorion membrane proteins in species of the melanogaster subgroup and found that chorion proteins as a group are indeed evolving faster than vitelline membrane proteins. At least one vitelline membrane protein (Vm32E), specifically localized on the outer eggshell, is also evolving faster than other vitelline membrane proteins suggesting that all proteins localized on the outer eggshell may be evolving rapidly. We also found evidence that specific codons in chorion proteins cp15 and cp16 are evolving under positive selection. Polymorphism surveys of cp16 revealed inflated levels of divergence relative to polymorphism in specific regions of the gene, indicating that these regions are under strong selection. At the morphological level, we found notable difference in eggshell surface morphologies between specialist (Drosophila sechellia and Drosophila erecta) and generalist species of Drosophila. We do not know if any of the chorion proteins actually interact with spermatozoids, therefore leaving the possibility of rapid evolution through gametic interaction wide open. At this point, however, our results support previous suggestions that divergences in ecology, particularly, ovipositional substrate divergences may be a strong force driving the evolution of eggshell proteins.     

Environmental heterogeneity,genotype-by-environment interactions and the reliability of sexual traits as indicators of mate quality

Exaggerated sexual displays are often supposed to indicate the indirect benefits females may receive from sexual reproduction with displaying males, but empirical evidence for positive relationships between the genetic quality and sexual trait quality is scant. The explanation for this might lie in the fact that mixing of reproductive individuals whose development has been influenced by genotype-by-environment interactions (GEIs) can blur the relationship between the individual male genetic quality and phenotype as perceived by females. Strong GEIs can generate an ecological crossover, where different genotypes are superior in environments that are separated either in space or time. Here, we use a stochastic simulation model to show that even a weak GEI, which does not generate an obvious ecological crossover, can neutralize or even reverse the relationship between genetic quality and sexual trait size in the presence of environmental heterogeneity during development. Our model highlights the importance of developmental selection in evolution of traits and allows us to predict the situations in which sexual displays might not be reliable indicators of genetic quality.     

Seasonal dynamics of entomopathogenic nematodes of the genera Steinernema and Heterorhabditis as a response to abiotic factors and abundance of insect hosts

The seasonal dynamics of entomopathogenic nematodes (EPNs) of the genus Steinernema and Heterorhabditis were studied during one season in meadow and oak wood habitats, in the vicinity of Ceské Budejovice, Czech Republic. The influences of soil temperature, moisture, and abundance of suitable hosts on EPN dynamics were investigated. The host range of these nematodes, in both habitats was also observed. A total of four EPN species were found in both habitats. Steinernema affine was the dominant species both in oak wood and in meadow. Additionally, the oak wood habitat was inhabited by S. kraussei and S. weiseri; the meadow habitat by Heterorhabditis bacteriophora. The mean abundance of total EPN community was 28,000ind./m(2) in oak wood and 11,000ind./m(2) in meadow. The seasonal dynamics of entomopathogenic nematodes in both habitats were characterized by high nematode densities in the beginning of the season, followed by a rapid decrease, and then stabilization. EPN abundances did not show any apparent correlation with soil temperature and moisture, but they were negatively correlated with the abundance of suitable insect hosts. Inter- and intraspecific competition for limited nutrients (hosts) probably played a major role in EPN seasonal dynamics. Broad host range of entomopathogenic nematodes in both habitats was predominantly represented by dipteran and coleopteran larvae. Most common hosts belonged to the families Asilidae, Bibionidae, and Empididae (Diptera), as well as Carabidae and Curculionidae (Coleoptera).     

Environmental heterogeneity influences the reliability of secondary sexual traits as condition indicators

Numerous studies have shown positive associations between ornaments and condition, as predicted by indicator models of sexual selection. However, this idea is continuously challenged by opposite results, which reveal our lack of full understanding of how sexual selection works. Environmental heterogeneity may explain such inconsistencies, but valid field tests of this idea are currently lacking. We first analysed the relationship between condition and ornament expression from nine populations over 7 years in a wild bird, the red grouse Lagopus lagopus scoticus. We then manipulated male aggressiveness at the population level by means of testosterone implants in a replicated field experiment. We found that the relationship between condition and ornamentation varied greatly between environments and became stronger when environmental conditions (ECs) were worse or when aggressiveness in the population was experimentally increased. Some ornaments may therefore reliably advertise a better condition only in adverse ECs. Considering environmental heterogeneity can help reconcile conflicting findings regarding the reliability of ornaments as indicators of condition and will help our understanding of sexual selection processes.     

Relative importance of the area and intensity of the orange spots of male guppies Poecilia reticulata as mating traits preferred by females

Digitally modified videos of male guppies Poecilia reticulata were used to examine the relative importance of the area and intensity of the orange spots as mating traits preferred by females. The females prioritized the area of the orange spots over intensity for their mate preference.     

Evolution of morphological differences with moderate genetic correlations among traits as exemplified by two flycatcher species (Ficeduia; Muscicapidae)

Theoretical work on multivariate evolution predicts that genetic correlations can act to constrain the rate at which new adaptive peaks are reached, but there is very limited empirical information available on this issue so far. To evaluate the importance of genetic correlations for evolutionary change, we studied the morphological differences between two flycatcher species (Ficeduia albicollis and F. hypoleuca) using both univariate and multivariate quantitative genetic models. Comparison of the results obtained using these different models revealed that even relatively low genetic correlations between traits will considerably increase the net selection forces needed for evolutionary changes in morphology. In particular, the divergence in wing and tail length, which are positively genetically correlated, would require a considerable amount of antagonistic selection. Because of the genetic correlations, strong selection will be needed to retain certain traits unchanged while others are changing. Based on these results, we argue that it is unlikely that small morphological differences such as between these two species could have evolved during a short (200 years) time period, i.e. the period of sympatry of these species in Sweden. These findings support the hypothesis that even relatively low genetic correlations may constrain short-term adaptive evolution in natural populations.     

The first proof of a possibility of change of temperature norms of insect development as a result of artificial selection for fast or slow development by the example of the red soldier bug <Emphasis Type="Italic">Pyrrhocoris apterus</Emphasis> (Heteroptera,Pyrrhocoridae)

For 3–4 generations, selection of the nymphs most rapidly and slowly developing at 28°C was performed in four families of the red soldier bug Pyrrhocoris apterus L. In each generation, duration of development of nymphs was determined at 5 constant temperatures from 20°C to 28°C. From these data the linear regression coefficient of the development rate against temperature (the development thermolability coefficient) for each generation was calculated. As a result of the selection the average duration of the nymph development was shortened or increased statistically significantly depending on its direction. The artificial selection for the development duration has been established to change not only this parameter, but also the temperature norms of the insect development. At selection for fast development the regression straight line slope (i.e., the regression coefficient value) increased statistically significantly, i.e., development became more dependent on temperature. As a rule, the temperature threshold of development increased. At selection for slow development the values of the regression coefficient and of the threshold decreased, but these differences were not statistically significant. The effect of artificial selection for the development duration on temperature norms of insect development has been revealed for the first time.