Good genes, genetic compatibility and the evolution of polyandry: use of the diallel cross to address competing hypotheses.

Genetic benefits can enhance the fitness of polyandrous females through the high intrinsic genetic quality of females' mates or through the interaction between female and male genes. I used a full diallel cross, a quantitative genetics design that involves all possible crosses among a set of genetically homogeneous lines, to determine the mechanism through which polyandrous female decorated crickets (Gryllodes sigillatus) obtain genetic benefits. I measured several traits related to fitness and partitioned the phenotypic variance into components representing the contribution of additive genetic variance ('good genes'), nonadditive genetic variance (genetic compatibility), as well as maternal and paternal effects. The results reveal a significant variance attributable to both nonadditive and additive sources in the measured traits, and their influence depended on which trait was considered. The lack of congruence in sources of phenotypic variance among these fitness-related traits suggests that the evolution and maintenance of polyandry are unlikely to have resulted from one selective influence, but rather are the result of the collective effects of a number of factors.

Ivy TM 《JOURNAL OF EVOLUTIONARY BIOLOGY》

The evolution of polyandry: intrinsic sire effects contribute to embryo viability.

Females typically mate with more than one male despite the costs incurred, thus questioning Bateman's principle. A series of genetic benefits have been proposed to account for the evolution of polyandry, including the acquisition of viability genes for offspring. The 'intrinsic male quality' hypothesis suggests that polyandry increases the probability that females produce offspring sired by males that bestow high viability on their offspring. Heritable variation in viability is the basic requirement for the occurrence of this genetic benefit. By using a half-sib breeding design with a species of cricket in which polyandry is known to increase hatching success, we present clear experimental evidence that intrinsic male quality contributes to embryo viability. Despite recent support for the evolution of polyandry based on compatibility of genotypes between males and females, we show that hatching success is not determined by an interaction between paternal and maternal genotypes but rather that sons inherit paternal genes that influence the viability of eggs laid by their mates. Moreover, our data implicate a potential role for indirect genetic effects of male accessory gland products on embryo viability. Additive genetic contributions to embryo viability may be an important factor underlying the frequently observed benefits of polyandrous behaviour.

García-González F ，Simmons LW 《JOURNAL OF EVOLUTIONARY BIOLOGY》

Maternal effects, but no good or compatible genes for sperm competitiveness in Australian crickets.

Explanations for the evolution of polyandry often center on the idea that females garner genetic benefits for their offspring by mating multiply. Furthermore, postcopulatory processes are thought to be fundamental to enabling polyandrous females to screen for genetic quality. Much attention has focused on the potential for polyandrous females to accrue such benefits via a sexy- or good-sperm mechanism, whereby additive variation exists among males in sperm competitiveness. Likewise, attention has focused on an alternative model, in which offspring quality (in this context, the sperm competitiveness of sons) hinges on an interaction between parental haplotypes (genetic compatibility). Sperm competitiveness that is contingent on parental compatibility will exhibit nonadditive genetic variation. We tested these models in the Australian cricket, Teleogryllus oceanicus, using a design that allowed us to partition additive, nonadditive genetic, and parental variance for sperm competitiveness. We found an absence of additive and nonadditive genetic variance in this species, challenging the direct relevance of either model to the evolution of sperm competitiveness in particular, and polyandry in general. Instead, we found maternal effects that were possibly sex-linked or cytoplasmically linked. We also found effects of focal male age on sperm competitiveness, with small increments in age conferring more competitive sperm.

Dowling DK ，Nystrand M ，Simmons LW 《-》

Sexual selection in the cricket Gryllus bimaculatus: no good genes?

Recent studies have suggested that females of the field cricket Gryllus bimaculatus exercise post-copulatory choice over the paternity of their offspring. There is evidence that these choices are made in relation to the genetic compatibility of mates rather than their absolute quality, but the magnitude of heritable differences in males has not been thoroughly examined. Using a half-sib breeding design we measured additive genetic variance and dam effects in a suite of reproductive and non-reproductive traits. Both components explained relatively little of the phenotypic variance across traits. The dam component in our design contains variance caused by both maternal effects and dominance. If maternal effects are negligible as suggested by previous studies, our data suggest that dominance variance is an important source of variation in these traits. The lack of additive genetic variation, but possible existence of large amounts of non-additive genetic variation is consistent with the idea that female mate choice and multiple mating may be driven by differences in genetic compatibility between potential mates rather than by differences in genetic quality.

Rodríguez-Muñoz R ，Bretman A ，Hadfield JD ，Tregenza T ... -  《GENETICA》

Analysis of the importance of genotypic variation, metabolic rate, morphology, sex and development time on immune function in the cricket, Gryllus firmus.

Immune defence is hypothesized to be a trait that bears significant fitness costs as well as benefits in that mounting a defence depreciates the value of other life-history traits. Thus the cost of mounting an immune response could affect the evolution of both the immune system and correlated life history traits. In this study we examined, by means of a diallel cross of four inbred lines, the genetic basis of two measures of immune function, metabolic rate and several traits in the sand cricket, Gryllus firmus. We specifically addressed the following questions: (1) is immune function determined primarily by genetic constitution or correlations with phenotypic traits that could reduce the effectiveness of the immune response; (2) do the two measures of immune function covary; (3) What are the contributions of additive, nonadditive and maternal effects to the immune function? As estimates of immune function, we used lytic activity and encapsulation rate. We found that inbred crickets were smaller than individuals from the crossed lines and took longer to develop. However, inbred lines did not differ from the crossed lines in immune function nor metabolic rates, suggesting that increased homozygosity has little or no effect on these traits in G. firmus. We found that both immune parameters showed significant genetic variation but no consistent relationships with the other phenotypic traits (metabolic rate, head width, body mass, development time and activity). There was significant additive genetic variation only in encapsulation rate, but, with the exception of the activity measure, significant nonadditive and reciprocal variances were found in all traits. Metabolic rate of crickets was heritable, but there was neither phenotypic nor genetic association between metabolic rate and the two parameters of immune function. Further, there was no correlation between these two measures. Females showed a higher encapsulation response than males, but there was no sex differences in lytic activity. Our study indicates that genetic variation in immune parameters can be a very significant contributor to phenotypic variation in immune function.

Rantala MJ ，Roff DA 《JOURNAL OF EVOLUTIONARY BIOLOGY》