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Biology subjects

Blows, M. W.

Publications and source records attributed to Blows, M. W..

5 recordsLinked to original sources

Using inbreeding to test the contribution of non-additive genetic effects to additive genetic variance: a case study in Drosophila serrata

Additive genetic variance, VA, is the key parameter for predicting adaptive and neutral phenotypic evolution. Changes in demography (e.g., increased close-relative inbreeding) can alter VA, but how depends on the, typically unknown, gene action and allele frequencies across many loci. For example, VA increases proportionally with the inbreeding coefficient when allelic effects are additive, but larger (or smaller) increases can occur when allele frequencies are unequal at causal loci with dominance effects. Here, we describe an experimental approach to assess the potential for rare, recessive alleles to inflate VA under inbreeding. Applying a powerful paired pedigree design in Drosophila serrata, we measured 11 wing traits on half-sibling families bred via either random or sibling mating, differing only in homozygosity (not allele frequency). Despite close inbreeding and substantial power to detect small VA, we detected no deviation from the expected additive effect of inbreeding on genetic (co)variances. Our results suggest the average dominance coefficient is very small relative to the additive effect, or that allele frequencies are relatively equal at loci affecting wing traits. We outline the further opportunities for this paired pedigree approach to reveal the characteristics of VA, providing insight into historical selection and future evolutionary potential.

evolutionary biology↗

Maintenance of quantitative genetic variance in complex, multi-trait phenotypes: The contribution of rare, large effect variants in two Drosophila species

The interaction of evolutionary processes to determine quantitative genetic variation has implications for contemporary and future phenotypic evolution, as well as for our ability to detect causal genetic variants. While theoretical studies have provided robust predictions to discriminate among competing models, empirical assessment of these has been limited. In particular, theory highlights the importance of pleiotropy in resolving observations of selection and mutation, but empirical investigations have typically been limited to few traits. Here, we applied high dimensional Bayesian Sparse Factor Genetic modelling to 3,385 gene expression traits from Drosophila melanogaster and from D. serrata to explore how genetic variance is distributed across high-dimensional phenotypic space. Surprisingly, most of the heritable trait covariation was due to few lines (genotypes) with extreme (>3 IQR from the median) values. This observation, in the two independently sampled species, suggests that the House of Cards (HoC) model might apply not only to individual expression traits, but also to emergent co-expression phenotypes. Intriguingly, while genotypes extreme for a multivariate factor also tended to have a higher proportion of individual traits that were extreme, we also observed genotypes that were outliers for multivariate factors but not for any individual traits. We observed other consistent differences between heritable multivariate factors with outlier lines versus those factors that conformed to a Gaussian distribution of genetic effects, including differences in gene functions. We use these observations to identify further data required to advance our understanding of the evolutionary dynamics and nature of standing genetic variation for quantitative traits.

genetics↗

Causes of variability in estimates of mutational variance from mutation accumulation experiments

Characteristics of the new phenotypic variation introduced via mutation have broad implications in evolutionary and medical genetics. Standardised estimates of this mutational variance, VM, span two orders of magnitude, but the causes of this remain poorly resolved. We investigated estimate heterogeneity using two approaches. First, meta-analyses of ~150 estimates from 37 mutation accumulation (MA) studies did not support a difference among taxa (which differ in mutation rate) in standardised VM, but provided equivocal support for standardised VM to vary with trait type (life history versus morphology, predicted to differ in mutation rate). Notably, several experimental factors were confounded with taxon and trait, and further empirical data are required to resolve their influences. Second, we analysed morphological data from an experiment in Drosophila serrata to determine the potential for unintentional heterogeneity among environments in which phenotypes were measured (i.e., among laboratories or time points) or transient segregation of mutations within MA lines to affect standardised VM. Approximating the size of an average MA experiment, variability among repeated estimates of (accumulated) mutational variance was comparable to variation among published estimates of standardised VM for morphological traits. This heterogeneity was (partially) attributable to unintended environmental variation or within line segregation of mutations only for wing size, not wing shape traits. We conclude that sampling error contributed substantial variation within this experiment, and infer that it will also contribute substantially to differences among published estimates. We suggest a logistically permissive approach to improve the precision of estimates, and consequently our understanding of the dynamics of mutational variance of quantitative traits.

genetics↗

Connecting micro and macroevolution using genetic incompatibilities and natural selection on additive genetic variance

Testing whether local adaptation and intrinsic reproductive isolation share a genetic basis can reveal important connections between adaptation and speciation. Local adaptation arises as advantageous alleles spread through a population, but whether these same advantageous alleles fail on the genetic backgrounds of other populations remains largely unknown. We used a quantitative genetic breeding design to produce a late generation (F4) recombinant hybrid population by equally mating amongst four contrasting ecotypes of a native Australian daisy for four generations. We tracked fitness across generations and measured morphological traits in the glasshouse, and used a reciprocal transplant to quantify fitness in all four parental habitats. In the glasshouse, plants in the second generation showed a reduction in fitness as a loss of fertility, but this was fully recovered in the following generation. The F4 hybrid lacked extreme phenotypes present in the parental ecotypes, suggesting that genes reducing hybrid fitness were linked to traits specific to each ecotype. In the natural habitats, additive genetic variance for fitness was greatest for habitats that showed stronger native-ecotype advantage, suggesting that a loss of genetic variation present in the parental ecotypes were adaptive in the natural habitats. Reductions in genetic variance for fitness were associated with a loss of ecological trade-offs previously described in the parental ecotypes. Furthermore, natural selection on morphological traits differed amongst the parental habitats, but was not predicted to occur towards the morphology of the parental ecotypes. Together, these results suggest that intrinsic reproductive isolation removed adaptive genetic variation present in the parental ecotypes. The evolution of these distinct ecotypes was likely governed by genetic variation that resulted in both ecological trade-offs and intrinsic reproductive isolation among populations adapted to contrasting environments.

evolutionary biology↗

Evolution of genetic variance during adaptive radiation

Genetic correlations between traits can bias adaptation away from optimal phenotypes and constrain the rate of evolution. If genetic correlations between traits limit adaptation to contrasting environments, rapid adaptive divergence across a heterogeneous landscape may be difficult. However, if genetic variance can evolve and align with the direction of natural selection, then abundant allelic variation can promote rapid divergence during adaptive radiation. Here, we explored adaptive divergence among ecotypes of an Australian native wildflower by quantifying divergence in multivariate phenotypes of populations that occupy four contrasting environments. We investigated differences in multivariate genetic variance underlying morphological traits and examined the alignment between divergence in phenotype and divergence in genetic variance. We found that divergence in mean multivariate phenotype has occurred along two major axes represented by different combinations of plant architecture and leaf traits. Ecotypes also showed divergence in the level of genetic variance in individual traits, and the multivariate distribution of genetic variance among traits. Divergence in multivariate phenotypic mean aligned with divergence in genetic variance, with most of the divergence in phenotype among ecotypes associated with a change in trait combinations that had substantial levels of genetic variance in each ecotype. Overall, our results suggest that divergent natural selection acting on high levels of standing genetic variation might fuel ecotypic differentiation during the early stages of adaptive radiation.

evolutionary biology↗