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1.  Global distribution of malaria-resistant MHC-HLA alleles: the number and frequencies of alleles and malaria risk 
Malaria Journal  2014;13(1):349.
Background
The major histocompatibility complex (MHC) is the most polymorphic genetic region in vertebrates, but the origin of such genetic diversity remains unresolved. Several studies have demonstrated at the within-population level that individuals harbouring particular alleles can be less or more susceptible to malaria, but these do not allow strong generalization.
Methods
Here worldwide data on the frequencies of several hundred MHC alleles of the human leucocyte antigen (HLA) system in relation to malaria risk at the between-population level were analysed in a phylogenetic framework, and results for different alleles were quantitatively summarized in a meta-analysis.
Results
There was an overall positive relationship between malaria pressure and the frequency of several HLA alleles indicating that allele frequencies increase in countries with strong malaria pressure. Nevertheless, considerable heterogeneity was observed across alleles, and some alleles showed a remarkable negative relationship with malaria risk. When heterogeneities were partitioned into different organization groups of the MHC, the strongest positive relationships were detected for alleles of the HLA-A and HLA-B loci, but there were also differences between MHC supertypes that constitute functionally distinct nucleotide sequences. Finally, the number of MHC alleles that are maintained within countries was also related to malaria risk.
Conclusion
Therefore, malaria represents a key selection pressure for the human MHC and has left clear evolutionary footprints on both the frequencies and the number of alleles observed in different countries.
Electronic supplementary material
The online version of this article (doi:10.1186/1475-2875-13-349) contains supplementary material, which is available to authorized users.
doi:10.1186/1475-2875-13-349
PMCID: PMC4162943  PMID: 25187124
Host-parasite interaction; Major histocompatibility complex; Malaria; Parasite transmission; Vector-borne infectious diseases
2.  Breeding Experience and the Heritability of Female Mate Choice in Collared Flycatchers 
PLoS ONE  2010;5(11):e13855.
Background
Heritability in mate preferences is assumed by models of sexual selection, and preference evolution may contribute to adaptation to changing environments. However, mate preference is difficult to measure in natural populations as detailed data on mate availability and mate sampling are usually missing. Often the only available information is the ornamentation of the actual mate. The single long-term quantitative genetic study of a wild population found low heritability in female mate ornamentation in Swedish collared flycatchers. One potentially important cause of low heritability in mate ornamentation at the population level is reduced mate preference expression among inexperienced individuals.
Methodology/Principal Findings
Applying animal model analyses to 21 years of data from a Hungarian collared flycatcher population, we found that additive genetic variance was 50 percent and significant for ornament expression in males, but less than 5 percent and non-significant for mate ornamentation treated as a female trait. Female breeding experience predicted breeding date and clutch size, but mate ornamentation and its variance components were unrelated to experience. Although we detected significant area and year effects on mate ornamentation, more than 85 percent of variance in this trait remained unexplained. Moreover, the effects of area and year on mate ornamentation were also highly positively correlated between inexperienced and experienced females, thereby acting to remove difference between the two groups.
Conclusions/Significance
The low heritability of mate ornamentation was apparently not explained by the presence of inexperienced individuals. Our results further indicate that the expression of mate ornamentation is dominated by temporal and spatial constraints and unmeasured background factors. Future studies should reduce unexplained variance or use alternative measures of mate preference. The heritability of mate preference in the wild remains a principal but unresolved question in evolutionary ecology.
doi:10.1371/journal.pone.0013855
PMCID: PMC2973971  PMID: 21079813
3.  Patterns of co-speciation and host switching in primate malaria parasites 
Malaria Journal  2009;8:110.
Background
The evolutionary history of many parasites is dependent on the evolution of their hosts, leading to an association between host and parasite phylogenies. However, frequent host switches across broad phylogenetic distances may weaken this close evolutionary link, especially when vectors are involved in parasites transmission, as is the case for malaria pathogens. Several studies suggested that the evolution of the primate-infective malaria lineages may be constrained by the phylogenetic relationships of their hosts, and that lateral switches between distantly related hosts may have been occurred. However, no systematic analysis has been quantified the degree of phylogenetic association between primates and their malaria parasites.
Methods
Here phylogenetic approaches have been used to discriminate statistically between events due to co-divergence, duplication, extinction and host switches that can potentially cause historical association between Plasmodium parasites and their primate hosts. A Bayesian reconstruction of parasite phylogeny based on genetic information for six genes served as basis for the analyses, which could account for uncertainties about the evolutionary hypotheses of malaria parasites.
Results
Related lineages of primate-infective Plasmodium tend to infect hosts within the same taxonomic family. Different analyses testing for congruence between host and parasite phylogenies unanimously revealed a significant association between the corresponding evolutionary trees. The most important factor that resulted in this association was host switching, but depending on the parasite phylogeny considered, co-speciation and duplication may have also played some additional role. Sorting seemed to be a relatively infrequent event, and can occur only under extreme co-evolutionary scenarios. The concordance between host and parasite phylogenies is heterogeneous: while the evolution of some malaria pathogens is strongly dependent on the phylogenetic history of their primate hosts, the congruent evolution is less emphasized for other parasite lineages (e.g. for human malaria parasites). Estimation of ancestral states of host use along the phylogenetic tree of parasites revealed that lateral transfers across distantly related hosts were likely to occur in several cases. Parasites cannot infect all available hosts, and they should preferentially infect hosts that provide a similar environment for reproduction. Marginally significant evidence suggested that there might be a consistent variation within host ranges in terms of physiology.
Conclusion
The evolution of primate malarias is constrained by the phylogenetic associations of their hosts. Some parasites can preserve a great flexibility to infect hosts across a large phylogenetic distance, thus host switching can be an important factor in mediating host ranges observed in nature. Due to this inherent flexibility and the potential exposure to various vectors, the emergence of new malaria disease in primates including humans cannot be predicted from the phylogeny of parasites.
doi:10.1186/1475-2875-8-110
PMCID: PMC2689253  PMID: 19463162
4.  Prevalence of avian influenza and host ecology 
Waterfowl and shorebirds are common reservoirs of the low pathogenic subtypes of avian influenza (LPAI), which are easily transmitted to poultry and become highly pathogenic. As the risk of virus transmission depends on the prevalence of LPAI in host-reservoir systems, there is an urgent need for understanding how host ecology, life history and behaviour can affect virus prevalence in the wild. To test for the most important ecological correlates of LPAI virus prevalence at the interspecific level, we applied a comparative analysis by using quantitative data on 30 bird species. We controlled for similarity among species due to common descent, differences in study effort and for covariance among ecological variables. We found that LPAI prevalence is a species-specific attribute and is a consequence of virus susceptibility, as it was negatively associated with the relative size of the bursa of Fabricius, an estimate of juvenile immune function. Species that migrate long distances have elevated prevalence of LPAI independent of phylogeny and other confounding factors. There was also a positive interspecific relationship between the frequency of surface feeding and virus prevalence, but this was sensitive to phylogenetic relatedness of species. Feeding in marine habitats is apparently associated with lower virus prevalence, but the effect of water salinity is likely to be indirect and affected by phylogeny. Our results imply that virus transmission via surface waters and frequent intra- and interspecific contacts during long migration are the major risk factors of avian influenza in the wild. However, the link between exploitation of surface waters and LPAI prevalence appears to be weaker than previously thought. This is the first interspecific study that provides statistical evidence that host ecology, immunity and phylogeny have important consequence for virus prevalence.
doi:10.1098/rspb.2007.0124
PMCID: PMC2275171  PMID: 17537707
Anatidae; bird flu; feeding ecology; migration; prevalence; risk assessment
5.  Birds Reveal their Personality when Singing 
PLoS ONE  2008;3(7):e2647.
Background
Individual differences in social behaviour may have consequences for mate choice and sexual signalling, because partners should develop preferences for personalities that maximize reproductive output. Here we propose that behavioural traits involved in sexual advertisement may serve as good indicators of personality, which is fundamental for sexual selection to operate on temperament. Bird song has a prominent and well-established role in sexual selection, and it displays considerable variation among individuals with a potentially strong personality component. Therefore, we predicted that features of song would correlate with estimates of personality.
Methodology/Principal Findings
In a field study of free-living male collared flycatchers, Ficedula albicollis, we characterised personality based on the exploration of an altered breeding environment, and based on the risk taken when a potential predator was approaching during a simulated territorial interaction. We found that explorative and risk-taker individuals consistently sang at lower song posts than shy individuals in the presence of a human observer. Moreover, males from lower posts established pair-bonds relatively faster than males from higher posts.
Conclusions/Significance
Our results may demonstrate that risk taking during singing correlates with risk taking during aggression and with exploration, thus personality may be manifested in different contexts involving sexual advertisement. These findings are in accordance with the hypothesis that the male's balance between investment in reproduction and risk taking is reflected in sexual displays, and it may be important information for choosy females that seek partners with personality traits enhancing breeding success.
doi:10.1371/journal.pone.0002647
PMCID: PMC2441454  PMID: 18612388
6.  Continental variation in relative hippocampal volume in birds: the phylogenetic extent of the effect and the potential role of winter temperatures 
Biology letters  2005;1(3):330-333.
Hippocampal (HC) volume has been hypothesized to increase with an increase in food-hoarding specialization in corvids and parids. Recent studies revealed that (i) the HC/hoarding relationship is significant when a difference in HC volume between Eurasian and North American species is controlled for and (ii) the evolutionary association has been acting on a broader phylogenetic context involving avian families outside the Corvidae and Paridae. However, the phylogenetic extent of the continent effect has not been previously addressed. Using data representing 48 avian species, we performed a phylogenetic analysis to test if continental effects are important in a wider evolutionary spectrum. Our results support the observation that Eurasian species have generally larger HC than North American species if variation in food hoarding, which also varied between continents, was held constant. Surprisingly, the relationship between continental distribution and relative HC volume was significant when we included only non-hoarding families in our analysis, indicating that the extent of the continent effect is much broader than originally described. We investigated the potential role of minimal winter temperatures at the northernmost distribution borders in mediating continent effects. The effect of winter temperatures on HC volume was weak and it did not vary consistently along continents. We suggest that the general continental differences in relative HC size are independent of food hoarding and that its determinants should be sought among other ecological factors and life-history traits.
doi:10.1098/rsbl.2005.0328
PMCID: PMC1523380  PMID: 16878181
Corvidae; food caching; hippocampus; Paridae; phylogeny
7.  Continental variation in relative hippocampal volume in birds: the phylogenetic extent of the effect and the potential role of winter temperatures 
Biology Letters  2005;1(3):330-333.
Hippocampal (HC) volume has been hypothesized to increase with an increase in food-hoarding specialization in corvids and parids. Recent studies revealed that (i) the HC/hoarding relationship is significant when a difference in HC volume between Eurasian and North American species is controlled for and (ii) the evolutionary association has been acting on a broader phylogenetic context involving avian families outside the Corvidae and Paridae. However, the phylogenetic extent of the continent effect has not been previously addressed. Using data representing 48 avian species, we performed a phylogenetic analysis to test if continental effects are important in a wider evolutionary spectrum. Our results support the observation that Eurasian species have generally larger HC than North American species if variation in food hoarding, which also varied between continents, was held constant. Surprisingly, the relationship between continental distribution and relative HC volume was significant when we included only non-hoarding families in our analysis, indicating that the extent of the continent effect is much broader than originally described. We investigated the potential role of minimal winter temperatures at the northernmost distribution borders in mediating continent effects. The effect of winter temperatures on HC volume was weak and it did not vary consistently along continents. We suggest that the general continental differences in relative HC size are independent of food hoarding and that its determinants should be sought among other ecological factors and life-history traits.
doi:10.1098/rsbl.2005.0328
PMCID: PMC1523380  PMID: 16878181
Corvidae; food caching; hippocampus; Paridae; phylogeny
8.  Sperm competition and sexually size dimorphic brains in birds 
Natural selection may favour sexually similar brain size owing to similar selection pressures in males and females, while sexual selection may lead to sexually dimorphic brains. For example, sperm competition involves clear-cut sex differences in behaviour, as males display, mate guard and copulate with females, while females choose among males, and solicit or reject copulations. These behaviours may require fundamentally different neural government in the two sexes leading to sex-dependent brain evolution. Using two phylogenetic approaches in a comparative study, we tested for roles of both natural and sexual-selection pressures on brain size evolution of birds. In accordance with the natural-selection theory, relative brain size of males coevolved with that of females, which may be the result of adaptation to similar environmental constraints such as feeding innovation. However, the mode of brain size evolution differed between the sexes, and factors associated with sperm competition as reflected by extra-pair paternity may give rise to sexually size dimorphic brains. Specifically, species in which females have larger brains than males were found to have a higher degree of extra-pair paternity independently of potentially confounding factors, whereas species in which males have relatively larger brains than females appeared to have lower rates of extra-pair paternity. Hence, the evolution of sperm competition may select for complex behaviours together with the associated neural substrates in the sex that has a higher potential to control extra-pair copulations at the observed levels. Brain function may thus be affected differently in males and females by sexual selection.
doi:10.1098/rspb.2004.2940
PMCID: PMC1634951  PMID: 15695206
birds; brain size; extra-pair paternity; phylogenetic correlation; sexual dimorphism
9.  Coevolving avian eye size and brain size in relation to prey capture and nocturnality. 
Behavioural adaptation to ecological conditions can lead to brain size evolution. Structures involved in behavioural visual information processing are expected to coevolve with enlargement of the brain. Because birds are mainly vision-oriented animals, we tested the predictions that adaptation to different foraging constraints can result in eye size evolution, and that species with large eyes have evolved large brains to cope with the increased amount of visual input. Using a comparative approach, we investigated the relationship between eye size and brain size, and the effect of prey capture technique and nocturnality on these traits. After controlling for allometric effects, there was a significant, positive correlation between relative brain size and relative eye size. Variation in relative eye and brain size were significantly and positively related to prey capture technique and nocturnality when a potentially confounding variable, aquatic feeding, was controlled statistically in multiple regression of independent linear contrasts. Applying a less robust, brunching approach, these patterns also emerged, with the exception that relative brain size did not vary with prey capture technique. Our findings suggest that relative eye size and brain size have coevolved in birds in response to nocturnal activity and, at least partly, to capture of mobile prey.
doi:10.1098/rspb.2002.1967
PMCID: PMC1690973  PMID: 12028780

Results 1-9 (9)