Reexamining Waddington: Canalization and new mutations are not required for the evolution of genetic assimilation

Over 65 years ago, Waddington demonstrated ancestrally phenotypically plastic traits can evolve to become constitutive, a process he termed genetic assimilation. Genetic assimilation evolves rapidly, assumed to be in large part due to segregating genetic variation only expressed in rare/novel environments, but otherwise phenotypically cryptic. Despite previous work suggesting a substantial role of cryptic genetic variation contributing to the evolution of genetic assimilation, some have argued for a prominent role for new mutations of large effect concurrent with selection. Interestingly, Waddington was less concerned by the relative contribution of CGV or new variants, but aimed to test the role of canalization, an evolved form of robustness. While canalization has been extensively studied, its role in the evolution of genetic assimilation is disputed, in part because explicit tests of evolved robustness are lacking. To address these questions, we recreated Waddington's selection experiments on an environmentally sensitive change in Drosophila wing morphology (crossvein development), using many independently evolved replicate lineages. Using these, we show that 1) a polygenic CGV, but not new variants of large effect are largely responsible for the evolved response demonstrated using both genomic and genetic approaches. 2) Using both environmental manipulations and mutagenesis of the evolved lineages that there is no evidence for evolved changes in canalization contributing to genetic assimilation. Finally, we demonstrate that 3) CGV has potentially pleiotropic and fitness consequences in natural populations and may not be entirely cryptic.

Intrapopulation Diversity of Chlamydomonas reinhardtii Response to Copper Ions: Growth and Photosynthetic Performance Under Stress

Despite being an essential micronutrient, copper is also a potentially toxic heavy metal. Using selection experiments, we produced Chlamydomonas reinhardtii populations with increased tolerance of copper ions and then derived pure cell lines from these populations. Strains derived from the same population (both adapted and nonadapted) significantly differed in terms of growth parameters. Cultivation of the strains in a range of copper ion concentrations revealed differences in growth and photosynthetic performance, which could be attributed to microevolutionary processes occurring with each cell division. Our results demonstrate the effects of environmental factors on rapidly multiplying microorganisms.

PSXIV-27 Simulation controlling inbreeding for selection experiments on woody breast meat in broiler

The purpose of this study was to control inbreeding for selection experiments on woody breast (WB) meat in broiler. The simulation was designed to figure out which mating plan would show proper breeding values while optimizing inbreeding assuming that a selection study would be done on WB in broilers starting with 500 males and 500 females as a foundational population. The simulated selections were based on Optimum Genetic Contribution theory (OGC) under different conditions over 10 generations, which uses relationships among individuals as weighting factors. It is selecting individuals by weighting estimated breeding values with average relationships among individuals. From the 2nd generation, various selection plans were considered in each sex, which were top 10, 20, 50 and 100 males selected, and top 100 and 200 females selected every generation. Each female bird was assumed to have 10 eggs. The algorithm is as follows: 1) Identify the individual having the best EBV; 2) Calculate average relationships between selected and candidates; 3) Select the individual having the best EBV adjusted for average relationships using the weighting factor k; 4) Repeat process until the number of individuals selected equals number required. Three different weighting values (k=0, 1, 2) were used, which made a total 24 different conditions compared (4×2×3). Additive genetic variance of breast meat was 1.134. Mendelian sampling terms were also considered when the breeding values were generated. Results showed that higher k value (k=2) controlled effectively inbreeding and maintained consistent increases in selection response. Differences in breeding values among selection plans with OGC algorithm and by EBV only was 4% on average; however, average rate of inbreeding (0.1) was controlled by 27% after 10 generations. These results indicate that the OGC algorithm can be used effectively in a short-term selection program with the relatively smaller number of populations.

Functional and Structural Segregation of Overlapping Helices in HIV-1

Overlapping coding regions balance selective forces between multiple genes. One possible division of nucleotide sequence is that the predominant selective force on a particular nucleotide can be attributed to just one gene. While this arrangement has been observed in regions in which one gene is structured and the other is disordered, we sought to explore how overlapping genes balance constraints when both protein products are structured over the same sequence. We use a combination of sequence analysis, functional assays and selection experiments to examine an overlapped region in HIV-1 that encodes helical regions in both Env and Rev. We find that functional segregation occurs even in this overlap, with each protein spacing its functional residues in a manner that allows a mutable non-binding face of one helix to encode important functional residues on a charged face in the other helix. Additionally, our experiments reveal novel and critical functional residues in Env and have implications for the therapeutic targeting of HIV-1.

Tell Me Where You’ve Been and I’ll Tell You How You’ll Evolve

ABSTRACT The reproducibility of adaptive evolution is a long-standing debate in evolutionary biology. Kempher et al. (M. L. Kempher, X. Tao, R. Song, B. Wu, et al., mBio 11:e00569-20, 2020, https://doi.org/10.1128/mBio.00569-20) used experimental evolution to investigate the effect of previous evolutionary trajectories on the ability of microbial populations to adapt to high temperatures. Despite the divergence caused by adaptation to previous environments, all populations reproducibly converged on similar final levels of fitness. Nevertheless, the genetic basis of adaptation depended on past selection experiments, reinforcing the idea that previous adaptation can dictate the trajectories of later evolutionary processes.

A combinatorial method to isolate short ribozymes from complex ribozyme libraries

In vitro selections are the only known methods to generate catalytic RNAs (ribozymes) that do not exist in nature. Such new ribozymes are used as biochemical tools, or to address questions on early stages of life. In both cases, it is helpful to identify the shortest possible ribozymes since they are easier to deploy as a tool, and because they are more likely to have emerged in a prebiotic environment. One of our previous selection experiments led to a library containing hundreds of different ribozyme clusters that catalyze the triphosphorylation of their 5′-terminus. This selection showed that RNA systems can use the prebiotically plausible molecule cyclic trimetaphosphate as an energy source. From this selected ribozyme library, the shortest ribozyme that was previously identified had a length of 67 nucleotides. Here we describe a combinatorial method to identify short ribozymes from libraries containing many ribozymes. Using this protocol on the library of triphosphorylation ribozymes, we identified a 17-nucleotide sequence motif embedded in a 44-nucleotide pseudoknot structure. The described combinatorial approach can be used to analyze libraries obtained by different in vitro selection experiments.

Estimating the genome-wide contribution of selection to temporal allele frequency change

Rapid phenotypic adaptation is often observed in natural populations and selection experiments. However, detecting the genome-wide impact of this selection is difficult since adaptation often proceeds from standing variation and selection on polygenic traits, both of which may leave faint genomic signals indistinguishable from a noisy background of genetic drift. One promising signal comes from the genome-wide covariance between allele frequency changes observable from temporal genomic data (e.g., evolve-and-resequence studies). These temporal covariances reflect how heritable fitness variation in the population leads changes in allele frequencies at one time point to be predictive of the changes at later time points, as alleles are indirectly selected due to remaining associations with selected alleles. Since genetic drift does not lead to temporal covariance, we can use these covariances to estimate what fraction of the variation in allele frequency change through time is driven by linked selection. Here, we reanalyze three selection experiments to quantify the effects of linked selection over short timescales using covariance among time points and across replicates. We estimate that at least 17 to 37% of allele frequency change is driven by selection in these experiments. Against this background of positive genome-wide temporal covariances, we also identify signals of negative temporal covariance corresponding to reversals in the direction of selection for a reasonable proportion of loci over the time course of a selection experiment. Overall, we find that in the three studies we analyzed, linked selection has a large impact on short-term allele frequency dynamics that is readily distinguishable from genetic drift.

Is there a genetic correlation between movement and immobility in field populations of a beetle?

Genetic correlations among behavioural traits are often controlled by pleiotropic genes. Many studies suggest the existence of genetic correlations among behavioural traits based on artificial selection experiments in the laboratory. However, few studies have examined whether behavioural correlations in the laboratory are maintained in the field, where natural selection works. Artificial selection experiments showed a behavioural correlation among death feigning, walking movement, and locomotor activity in the red flour beetle (Tribolium castaneum). This study investigated whether this behavioural correlation is observed in wild T. castaneum populations. We also collected beetles from various regions in Japan and investigated the geographic variation in these traits. There was geographic variation in the three behavioural traits. However, these behavioural traits were not correlated. The results suggest that the genetic correlations among behavioural traits are not maintained in the field. Therefore, the results derived from laboratory experiments may be overestimated. The same correlation between traits was not believed to arise in the field, as the indoor results may have been caused by unrealistic selection pressures. Further laboratory and field investigations are both needed.