Protein deprivation decreases male survival and the intensity of sexual antagonism in southern field crickets Gryllus bimaculatus.

Recent theory predicts that the magnitude of sexual antagonism should depend on how well populations are adapted to their environment. We tested this idea experimentally by comparing intersexual genetic correlations for adult survival in pedigreed populations of southern field crickets (Gryllus bimaculatus) raised on naturally balanced (free-choice) vs. imbalanced (protein-deprived) diets. We tested for (1) sex differences in nutritional intake and preference, (2) sex-specific effects of protein deprivation on survival and (3) diet dependence of the level of sexual antagonism. Adult males and females consumed a similar amount of protein, but protein deprivation decreased male survival but not female survival. Protein deprivation appeared to decrease the degree of sexual antagonism as intersexual genetic correlations were significantly lower than 1 only for the complementary free-choice diet group but close to 1 for the protein-deficient diet group. Our findings thereby implied that variation in nutritional environments can alter the magnitude of sexual antagonism. This research represents an important step towards understanding the relationship between sexual antagonism and adaptation in heterogeneous environments.

The correlation between coloration and exploration behaviour varies across hierarchical levels in a wild passerine bird.

In vertebrates, darker individuals are often found to be more active and willing to take risks (representing characteristics of a 'proactive' coping style), whereas lighter individuals are instead more cautious and less active (representing characteristics of a 'reactive' coping style). It is thus generally expected that melanin-based coloration and proactivity form a suite of positively integrated traits at the among-individual level. Here, we use a multigenerational pedigree of free-living great tits (Parus major) to partition variation in, and the correlation between, melanin-based breast stripe ('tie') size and exploration behaviour (a proxy for coping style) into its among- and within-individual components. We show that both traits harbour heritable variation. Against predictions, tie size and speed of exploration were negatively correlated at the among-individual level due to the combined influences of permanent environmental and additive genetic effects. By contrast, the two traits were weakly positively correlated within individuals (i.e. individuals increasing in tie size after moult tended to become more explorative). The patterns of among-individual covariance were not caused by correlational selection as we found additive and opposite selection pressures acting on the two traits. These findings imply that testing hypotheses regarding the existence of a 'syndrome' at the among-individual level strictly requires variance partitioning to avoid inappropriate interpretations as the negative 'unpartitioned' phenotypic correlation between exploration and tie size resulted from counteracting effects of within- and among-individual correlations. Identifying sources and levels of (co)variation in phenotypic traits is thus critical to our understanding of biological patterns and evolutionary processes.

Behaviour-related DRD4 polymorphisms in invasive bird populations.

It has been suggested that individual behavioural traits influence the potential to successfully colonize new areas. Identifying the genetic basis of behavioural variation in invasive species thus represents an important step towards understanding the evolutionary potential of the invader. Here, we sequenced a candidate region for neophilic/neophobic and activity behaviour - the complete exon 3 of the DRD4 gene - in 100 Yellow-crowned bishops (Euplectes afer) from two invasive populations in Spain and Portugal. The same birds were scored twice for activity behaviour while exposed to novel objects (battery or slice of apple) in captivity. Response to novel objects was repeatable (r = 0.41) within individuals. We identified two synonymous DRD4 SNPs that explained on average between 11% and 15% of the phenotypic variance in both populations, indicating a clear genetic component to the neophilic/neophobic/activity personality axis in this species. This consistently high estimated effect size was mainly due to the repeated measurement design, which excludes part of the within-individual nongenetic variance in the response to different novel objects. We suggest that the alternative alleles of these SNPs are likely introduced from the original population and maintained by weak or antagonistic selection during different stages of the invasion process. The identified genetic variants have not only the potential to serve as genetic markers of the neophobic/neophilic/activity personality axis, but may also help to understand the evolution of behaviour in these invasive bird populations.

Exploring patterns of variation in clutch size-density reaction norms in a wild passerine bird.

Negative density dependence of clutch size is a ubiquitous characteristic of avian populations and is partly due to within-individual phenotypic plasticity. Yet, very little is known about the extent to which individuals differ in their degree of phenotypic plasticity, whether such variation has a genetic basis and whether level of plasticity can thus evolve in response to selection. Using 18 years of data of a Dutch great tit population (Parus major), we show that females reduced clutch size with increasing population density (slopes of the reaction norms), differed strongly in their average clutch size (elevations of the reaction norms) at the population-mean density and that the latter variation was partly heritable. In contrast, we could not detect individual variation in phenotypic plasticity ('I × E'). Level of plasticity is thus not likely to evolve in response to selection in this population. Observed clutch sizes deviated more from the estimated individual reaction norms in certain years and densities, implying that the within-individual between-year variance (so-called residual variance) of clutch size was heterogeneous with respect to these factors. Given the observational nature of this study, experimental manipulation of density is now warranted to confirm the causality of the observed density effects. Our analyses demonstrate that failure to acknowledge this heterogeneity would have inflated the estimate of 'I × E' and led to misinterpretation of the data. This paper thereby emphasizes the fact that heterogeneity in residuals can provide biologically insightful information about the ecological processes underlying the data.

Parental provisioning behaviour plays a key role in linking personality with reproductive success.

Repeatable behavioural traits ('personality') have been shown to covary with fitness, but it remains poorly understood how such behaviour-fitness relationships come about. We applied a multivariate approach to reveal the mechanistic pathways by which variation in exploratory and aggressive behaviour is translated into variation in reproductive success in a natural population of blue tits, Cyanistes caeruleus. Using path analysis, we demonstrate a key role for provisioning behaviour in mediating the link between personality and reproductive success (number of fledged offspring). Aggressive males fed their nestlings at lower rates than less aggressive individuals. At the same time, their low parental investment was associated with increased female effort, thereby positively affecting fledgling production. Whereas male exploratory behaviour was unrelated to provisioning behaviour and reproductive success, fast-exploring females fed their offspring at higher rates and initiated breeding earlier, thus increasing reproductive success. Our findings provide strong support for specific mechanistic pathways linking components of behavioural syndromes to reproductive success. Importantly, relationships between behavioural phenotypes and reproductive success were obscured when considering simple bivariate relationships, underlining the importance of adopting multivariate views and statistical tools as path analysis to the study of behavioural evolution.

Quantitative genetics of behavioural reaction norms: genetic correlations between personality and behavioural plasticity vary across stickleback populations.

Behavioural ecologists have proposed various evolutionary mechanisms as to why different personality types coexist. Our ability to understand the evolutionary trajectories of personality traits requires insights from the quantitative genetics of behavioural reaction norms. We assayed > 1000 pedigreed stickleback for initial exploration behaviour of a novel environment, and subsequent changes in exploration over a few hours, representing their capacity to adjust their behaviour to changes in perceived novelty and risk. We found heritable variation in both the average level of exploration and behavioural plasticity, and population differences in the sign of the genetic correlation between these two reaction norm components. The phenotypic correlation was not a good indicator of the genetic correlation, implying that quantitative genetics are necessary to appropriately evaluate evolutionary hypotheses in cases such as these. Our findings therefore have important implications for future studies concerning the evolution of personality and plasticity.

Early exposure to nonlethal predation risk by size-selective predators increases somatic growth and decreases size at adulthood in three-spined sticklebacks.

Predation has an important influence on life history traits in many organisms, especially when they are young. When cues of trout were present, juvenile sticklebacks grew faster. The increase in body size as a result of exposure to cues of predators was adaptive because larger individuals were more likely to survive predation. However, sticklebacks that had been exposed to cues of predators were smaller at adulthood. This result is consistent with some life history theory. However, these results prompt an alternative hypothesis, which is that the decreased size at adulthood reflects a deferred cost of early rapid growth. Compared to males, females were more likely to survive predation, but female size at adulthood was more affected by cues of predators than male size at adulthood, suggesting that size at adulthood might be more important to male fitness than to female fitness.