Markus J. Rantala

Do pheromones reveal male immunocompetence

Pheromones function not only as mate attractors, but they may also relay important information to prospective mates. It has been shown that vertebrates can distinguish, via olfactory mechanisms, major histocompatibility complex types in their prospective mates. However, whether pheromones can transmit information about immunocompetence is unknown. Here, we show that female mealworm beetles (Tenebrio molitor) prefer pheromones from males with better immunocompetence, indicated by a faster encapsulation rate against a novel antigen, and higher levels of phenoloxidase in haemolymph. Thus, the present study indicates that pheromones could transmit information about males parasite resistance ability and may work as a reliable sexual ornament for female choice.

Interspecific aggression and character displacement in the damselfly Calopteryx splendens.

Problems in species recognition are thought to affect the evolution of secondary sexual characters mainly through avoidance of maladaptive hybridization. Another, but much less studied avenue for the evolution of sexual characters due to species recognition problems is through interspecific aggression. In the damselfly, Calopteryx splendens, males have pigmented wing spots as a sexual character. Large‐spotted males resemble males of another species, Calopteryx virgo, causing potential problems in species recognition. In this study, we investigate whether there is character displacement in wing spot size and whether interspecific aggression could cause this pattern. We found first that wing spot size of C. splendens in populations decreased with increasing relative abundance of C. virgo. Secondly, C. virgo males were more aggressive towards large‐ than small‐spotted C. splendens males. Thirdly, in interspecific contests C. virgo males had better territory holding ability than C. splendens males. These results suggest that interspecific aggression may have caused character displacement in wing spot size of C. splendens, because the intensity of aggression towards large‐spotted males is likely to increase with relative abundance of C. virgo males. Thus, interspecific aggression may be an evolutionarily significant force that is able to cause divergence in secondary sexual characters.

Effect of bacterial infection on antioxidant activity and lipid peroxidation in the midgut of Galleria mellonella L. larvae (Lepidoptera, Pyralidae)

Bacillus thuringiensis is one of the most widely used sources of biorational pesticides, as well as a key source of genes for transgenic expression to provide pest resistance in plants. In this study the effect of Bacillus thuringiensis ssp. galleriae (Bt) infection on the activity of superoxide dismutase (SOD), glutathione S-transferase (GST), catalase (CAT), concentrations of oxidated and reduced thiols (RSSR/RSH) and malondialdehyde (MDA) was tested in the midgut of Galleria mellonella larvae. We found that Bt infection resulted in increased activities of SOD, GST, malondialdehyde and RSSR/RSH ratio the first day after inoculation. However, catalase activity decreased on the first and following days after bacterial infection by Bt. Our results confirm the hypothesis that Bt infection increases the level of oxidative stress in the larval midgut. In light of this study, it seems possible that oxidative damage contributes to cell death in the midgut during bacteriosis.

Natural host-plant quality affects immune defence of an insect herbivore

We examined the effect of natural host‐plant quality on immune resistance in the autumnal moth, Epirrita autumnata (Borkhausen) (Lepidoptera: Geometridae). The division of mountain birches [Betula pubescens ssp. czerepanovii (Orlova) Hämet‐ahti (Betulaceae)] into two categories, high‐ and low‐quality food for larvae, was based on previous years’ results on the relative growth rate of the autumnal moth on the trees selected. The strength of the immune defence of autumnal moths was determined by measuring their encapsulation rate to exposure to a foreign antigen and the phenoloxidase (PO) activity of the pupal haemolymph. We found that individuals reared as larvae on naturally low‐quality food had a significantly higher encapsulation rate at the pupal stage than individuals reared on high‐quality food. Females also had a higher encapsulation rate than males. Food quality did not have statistically significant effect on PO activity, nor did this response variable show any differences between the sexes. Using half‐sib analyses, we found significant heritable variation in the encapsulation rate; the heritable variation in PO activity was near to significant, although equally strong. Heritability estimates (h2: 0.19–0.27) for immune defence traits were relatively low and only moderate when compared to other studies with insects. We also found a negative genetic correlation between pupal mass and PO activity, but not between PO activity and encapsulation rate. Our results suggest that the quality of food affects immune defence in the autumnal moth. Thus an intricate tritrophic relationship exists between the folivorous insect, the host tree, and the insects natural enemies (e.g., pathogens, parasites, and parasitoids). This study demonstrates that natural variation within a food plant species has an effect on the innate immune system of an herbivorous insect.

A trade-off between sexual signalling and immune function in a natural population of the drumming wolf spider Hygrolycosa rubrofasciata.

The field of ecological immunology is ultimately seeking to address the question ‘Why is there variation in immune function?’ Here, we provide experimental evidence that costs of ubiquitous sexual signals are a significant source of variation in immune function. In the mating season, males of the wolf spider Hygrolycosa rubrofasciata drum against dry leaves while wandering around the habitat searching for receptive females. According to a previous study, the male metabolic rate during the drumming increases 22‐fold compared to the resting metabolic rate. In the present study, we examined whether investment in costly courtship drumming decreases male immune function in a wild population of H. rubrofasciata. We induced males to increase their drumming rate by introducing females in proximity. As estimates of male immune function, we used lytic activity and encapsulation rate. Lytic activity estimates the concentration of antimicrobial peptides in haemolymph, which have been shown to play an important role in defence against bacteria, viruses and fungi. Encapsulation is an important defence mechanism against nematodes and insect parasitoids, but it also plays a role in defence against viruses. Our results show that males with nonarbitrarily increased investment in drumming rate had considerably lower lytic activities than control males. Also, there was a tendency for males with nonarbitrarily increased investment in drumming rate to have lower encapsulation rates than control males. This study provides experimental evidence for the first time, to our knowledge, that there are direct immunological costs of sexual signalling in natural populations. Therefore, immunological costs of sexual signals may provide significant phenotypic variation to parasite‐mediated sexual selection.

DEFOLIATING INSECT IMMUNE DEFENSE INTERACTS WITH INDUCED PLANT DEFENSE DURING A POPULATION OUTBREAK

During population outbreaks, top-down and bottom-up factors are unable to control defoliator numbers. To our knowledge, details of biotic interactions leading to increased population density have not been studied during real population outbreaks. We experimentally assessed the strength of plant defenses and of insect immunocompetence, assumed to contribute to active insect resistance against parasitoids and pathogens, in the geometrid Epirrita autumnata during a steep increase in population density. We demonstrated rapid (same-season) induced resistance in the foliage of its host, mountain birch. The response was systemic, spreading throughout the tree, and retarded larval growth rate by approximately 10%. On the other hand, no direct delayed carry-over effects were found in the next season in larval growth rate, mortality, or pupal mass. Larval damage to a tree during the previous year, however, significantly (by approximately 13%) accelerated the advance of the immune response (measured as melanization of an implant inserted into the pupal hemocoel). The encapsulation rate correlated positively with larval mortality in trees in which larvae had been introduced the previous year, but not in control trees. Both of these observations suggest that induced plant defense was associated with an increased insect immunocompetence during the population increase.

Starvation Reveals Maintenance Cost of Humoral Immunity

Susceptibility to pathogens and genetic variation in disease resistance is assumed to persist in nature because of the high costs of immunity. Within immunity there are different kinds of costs. Costs of immunological deployment, the costs of mounting an immune response, are measured as a change in fitness following immunological challenge. Maintenance costs of immunity are associated with investments of resources into the infrastructure of an immune system and keeping the system at a given level of readiness in the absence of infection. To demonstrate the costs of immunological maintenance in the absence of infection is considered more difficult. In the present study we examined the maintenance costs of the immune system in lines of Drosophila melanogaster that differed in their antibacterial innate immune response under starved and non-starved conditions. Immunodeficient mutant flies that have to invest less in the immunological maintenance were found to live longer under starvation than wild type flies, whereas the opposite was found when food was provided ad libitum. Our study provides evidence for the physiological cost of immunological maintenance and highlights the importance of environmental variation in the study of evolutionary trade-offs.

Foliar Phenolics are Differently Associated with Epirrita autumnata Growth and Immunocompetence

The quality of available food may affect insect herbivores directly (via growth and survivorship) and/or indirectly (by modifying insect vulnerability to parasitoids and pathogens). We examined the relationship between different phenolic compounds, belonging to various phenolic groups, in Betula pubescens spp. czerepanovii (mountain birch) foliage and the larval performance of the geometrid Epirrita autumnata (autumnal moth). Direct effects on insect performance were described by pupal weight, developmental rate, and survivorship; indirect effects were described by the encapsulation rate of an implant inserted into the insect hemocoel, a commonly used way to describe insect immune defense. We found profound differences in the effects of different phenolic categories: several individual hydrolyzable tannins were associated positively with larval performance but negatively with level of immune defense, whereas flavonoid glycosides were inversely related to larval survival but showed no association with the larvae immune defense.

Adiposity, compared with masculinity, serves as a more valid cue to immunocompetence in human mate choice

According to the ‘good genes’ hypothesis, females choose males based on traits that indicate the males genetic quality in terms of disease resistance. The ‘immunocompetence handicap hypothesis’ proposed that secondary sexual traits serve as indicators of male genetic quality, because they indicate that males can contend with the immunosuppressive effects of testosterone. Masculinity is commonly assumed to serve as such a secondary sexual trait. Yet, women do not consistently prefer masculine looking men, nor is masculinity consistently related to health across studies. Here, we show that adiposity, but not masculinity, significantly mediates the relationship between a direct measure of immune response (hepatitis B antibody response) and attractiveness for both body and facial measurements. In addition, we show that circulating testosterone is more closely associated with adiposity than masculinity. These findings indicate that adiposity, compared with masculinity, serves as a more important cue to immunocompetence in female mate choice.

Transgenerational effects of parental larval diet on offspring development time, adult body size and pathogen resistance in Drosophila melanogaster.

Environmental conditions experienced by parents are increasingly recognized to affect offspring performance. We set out to investigate the effect of parental larval diet on offspring development time, adult body size and adult resistance to the bacterium Serratia marcescens in Drosophila melanogaster. Flies for the parental generation were raised on either poor or standard diet and then mated in the four possible sex-by-parental diet crosses. Females that were raised on poor food produced larger offspring than females that were raised on standard food. Furthermore, male progeny sired by fathers that were raised on poor food were larger than male progeny sired by males raised on standard food. Development times were shortest for offspring whose one parent (mother or the father) was raised on standard and the other parent on poor food and longest for offspring whose parents both were raised on poor food. No evidence for transgenerational effects of parental diet on offspring disease resistance was found. Although paternal effects have been previously demonstrated in D. melanogaster, no earlier studies have investigated male-mediated transgenerational effects of diet in this species. The results highlight the importance of not only considering the relative contribution each parental sex has on progeny performance but also the combined effects that the two sexes may have on offspring performance.