Compensatory trans‐regulatory alleles minimizing variation in TDH3 expression are common within Saccharomyces cerevisiae

Heritable variation in gene expression is common within species. Much of this variation is due to genetic differences outside of the gene with altered expression and is trans‐acting. This trans‐regulatory variation is often polygenic, with individual variants typically having small effects, making the genetic architecture and evolution of trans‐regulatory variation challenging to study. Consequently, key questions about trans‐regulatory variation remain, including the variability of trans‐regulatory variation within a species, how selection affects trans‐regulatory variation, and how trans‐regulatory variants are distributed throughout the genome and within a species. To address these questions, we isolated and measured trans‐regulatory differences affecting TDH3 promoter activity among 56 strains of Saccharomyces cerevisiae, finding that trans‐regulatory backgrounds varied approximately twofold in their effects on TDH3 promoter activity. Comparing this variation to neutral models of trans‐regulatory evolution based on empirical measures of mutational effects revealed that despite this variability in the effects of trans‐regulatory backgrounds, stabilizing selection has constrained trans‐regulatory differences within this species. Using a powerful quantitative trait locus mapping method, we identified ∼100 trans‐acting expression quantitative trait locus in each of three crosses to a common reference strain, indicating that regulatory variation is more polygenic than previous studies have suggested. Loci altering expression were located throughout the genome, and many loci were strain specific. This distribution and prevalence of alleles is consistent with recent theories about the genetic architecture of complex traits. In all mapping experiments, the nonreference strain alleles increased and decreased TDH3 promoter activity with similar frequencies, suggesting that stabilizing selection maintained many trans‐acting variants with opposing effects. This variation may provide the raw material for compensatory evolution and larger scale regulatory rewiring observed in developmental systems drift among species.Peer Reviewedhttps://deepblue.lib.umich.edu/bitstream/2027.42/151914/1/evl3137_am.pdfhttps://deepblue.lib.umich.edu/bitstream/2027.42/151914/2/evl3137.pd

Genetic basis of octanoic acid resistance in Drosophila sechellia: functional analysis of a fine‐mapped region

Drosophila sechellia is a species of fruit fly endemic to the Seychelles islands. Unlike its generalist sister species, D. sechellia has evolved to be a specialist on the host plant Morinda citrifolia. This specialization is interesting because the plant’s fruit contains secondary defence compounds, primarily octanoic acid (OA), that are lethal to most other Drosophilids. Although ecological and behavioural adaptations to this toxic fruit are known, the genetic basis for evolutionary changes in OA resistance is not. Prior work showed that a genomic region on chromosome 3R containing 18 genes has the greatest contribution to differences in OA resistance between D. sechellia and D. simulans. To determine which gene(s) in this region might be involved in the evolutionary change in OA resistance, we knocked down expression of each gene in this region in D. melanogaster with RNA interference (RNAi) (i) ubiquitously throughout development, (ii) during only the adult stage and (iii) within specific tissues. We identified three neighbouring genes in the Osiris family, Osiris 6 (Osi6), Osi7 and Osi8, that led to decreased OA resistance when ubiquitously knocked down. Tissue‐specific RNAi, however, showed that decreasing expression of Osi6 and Osi7 specifically in the fat body and/or salivary glands increased OA resistance. Gene expression analyses of Osi6 and Osi7 revealed that while standing levels of expression are higher in D. sechellia, Osi6 expression is significantly downregulated in salivary glands in response to OA exposure, suggesting that evolved tissue‐specific environmental plasticity of Osi6 expression may be responsible for OA resistance in D. sechellia.Peer Reviewedhttps://deepblue.lib.umich.edu/bitstream/2027.42/136293/1/mec14001_am.pdfhttps://deepblue.lib.umich.edu/bitstream/2027.42/136293/2/mec14001.pdfhttps://deepblue.lib.umich.edu/bitstream/2027.42/136293/3/mec14001-sup-0001-SupInfo.pd

Evolution of the Spider Homeobox Gene Repertoire by Tandem and Whole Genome Duplication

Gene duplication generates new genetic material that can contribute to the evolution of gene regulatory networks and phenotypes. Duplicated genes can undergo subfunctionalization to partition ancestral functions and/or neofunctionalization to assume a new function. We previously found there had been a whole genome duplication (WGD) in an ancestor of arachnopulmonates, the lineage including spiders and scorpions but excluding other arachnids like mites, ticks, and harvestmen. This WGD was evidenced by many duplicated homeobox genes, including two Hox clusters, in spiders. However, it was unclear which homeobox paralogues originated by WGD versus smaller-scale events such as tandem duplications. Understanding this is a key to determining the contribution of the WGD to arachnopulmonate genome evolution. Here we characterized the distribution of duplicated homeobox genes across eight chromosome-level spider genomes. We found that most duplicated homeobox genes in spiders are consistent with an origin by WGD. We also found two copies of conserved homeobox gene clusters, including the Hox, NK, HRO, Irx, and SINE clusters, in all eight species. Consistently, we observed one copy of each cluster was degenerated in terms of gene content and organization while the other remained more intact. Focussing on the NK cluster, we found evidence for regulatory subfunctionalization between the duplicated NK genes in the spider Parasteatoda tepidariorum compared to their single-copy orthologues in the harvestman Phalangium opilio. Our study provides new insights into the relative contributions of multiple modes of duplication to the homeobox gene repertoire during the evolution of spiders and the function of NK genes

Fitness effects of altering gene expression noise in <i>Saccharomyces cerevisiae</i>

Gene expression noise is an evolvable property of biological systems that describes differences in expression among genetically identical cells in the same environment. Prior work has shown that expression noise is heritable and can be shaped by selection, but the impact of variation in expression noise on organismal fitness has proven difficult to measure. Here, we quantify the fitness effects of altering expression noise for the TDH3 gene in Saccharomyces cerevisiae. We show that increases in expression noise can be deleterious or beneficial depending on the difference between the average expression level of a genotype and the expression level maximizing fitness. We also show that a simple model relating single-cell expression levels to population growth produces patterns consistent with our empirical data. We use this model to explore a broad range of average expression levels and expression noise, providing additional insight into the fitness effects of variation in expression noise

Recurrent Modification of a Conserved Cis-Regulatory Element Underlies Fruit Fly Pigmentation Diversity

The development of morphological traits occurs through the collective action of networks of genes connected at the level of gene expression. As any node in a network may be a target of evolutionary change, the recurrent targeting of the same node would indicate that the path of evolution is biased for the relevant trait and network. Although examples of parallel evolution have implicated recurrent modification of the same gene and cis-regulatory element (CRE), little is known about the mutational and molecular paths of parallel CRE evolution. In Drosophila melanogaster fruit flies, the Bric-à-brac (Bab) transcription factors control the development of a suite of sexually dimorphic traits on the posterior abdomen. Female-specific Bab expression is regulated by the dimorphic element, a CRE that possesses direct inputs from body plan (ABD-B) and sex-determination (DSX) transcription factors. Here, we find that the recurrent evolutionary modification of this CRE underlies both intraspecific and interspecific variation in female pigmentation in the melanogaster species group. By reconstructing the sequence and regulatory activity of the ancestral Drosophila melanogaster dimorphic element, we demonstrate that a handful of mutations were sufficient to create independent CRE alleles with differing activities. Moreover, intraspecific and interspecific dimorphic element evolution proceeded with little to no alterations to the known body plan and sex-determination regulatory linkages. Collectively, our findings represent an example where the paths of evolution appear biased to a specific CRE, and drastic changes in function were accompanied by deep conservation of key regulatory linkages. © 2013 Rogers et al

Theta-point behavior of diluted polymer solutions: Can one observe the universal logarithmic corrections predicted by field theory?

In recent large scale Monte-Carlo simulations of various models of Theta-point polymers in three dimensions Grassberger and Hegger found logarithmic corrections to mean field theory with amplitudes much larger than the universal amplitudes of the leading logarithmic corrections calculated by Duplantier in the framework of tricritical O(n) field theory. To resolve this issue we calculate the universal subleading correction of field theory, which turns out to be of the same order of magnitude as the leading correction for all chain lengths available in present days simulations. Borel resummation of the renormalization group flow equations also shows the presence of such large corrections. This suggests that the published simulations did not reach the asymptotic regime. To further support this view, we present results of Monte-Carlo simulations on a Domb-Joyce like model of weakly interacting random walks. Again the results cannot be explained by keeping only the leading corrections, but are in fair accord with our full theoretical result. The corrections found for the Domb-Joyce model are much smaller than those for other models, which clearly shows that the effective corrections are not yet in the asymptotic regime. All together our findings show that the existing simulations of Theta-polymers are compatible with tricritical field theory since the crossover to the asymptotic regime is very slow. Similar results were found earlier for self avoiding walks at their upper critical dimension d=4.Comment: 15 pages,6 figure

Bighorn Basin Coring Project (BBCP): a continental perspective on early Paleogene hyperthermals

During the summer of 2011, the Bighorn Basin Coring Project (BBCP) recovered over 900m of overlapping core from 3 different sites in late Paleocene to early Eocene fluvial deposits of northwestern Wyoming. BBCP cores are being used to develop high-resolution proxy records of the Paleocene–Eocene Thermal Maximum (PETM) and Eocene Thermal Maximum 2 (ETM2) hyperthermal events. These events are short-term, large magnitude global warming events associated with extreme perturbations to the earth’s carbon cycle. Although the PETM and ETM2 occurred ~55–52 million years ago, they are analogous in many ways to modern anthropogenic changes to the carbon cycle. By applying various sedimentological, geochemical, and palynological methods to the cores, we hope to better understand what caused these events, study the biogeochemical and ecological feedbacks that operated during them, and reveal precisely how they impacted continental environments. Core recovery was > 98% in all holes and most drilling was carried out without fluid additives, showing that continuous coring of continental smectitic deposits like these can be achieved with minimal risk of contamination to molecular biomarkers. Cores were processed in the Bremen Core Repository where the science team convened for 17 days to carry out data collection and sampling protocols similar to IODP projects. Initial results show that the weathered horizon extends to as much as ~30m below the surface and variations in magnetic susceptibility within the cores record an interplay between grain size and pedogenesis. Previous investigations of outcrops near the BBCP drill sites allow detailed evaluation of the effects of weathering on common proxy methods. Studies of lithofacies, organic geochemistry, stable isotope geochemistry, calibrated XRF core scanning, paleomagnetics, and palynology are underway and will represent the highest resolution and most integrated proxy records of the PETM from a continental setting yet known. An extensive outreach program is in place to capitalize on the educational value associated with the Bighorn Basin’s unusually complete record of Phanerozoic earth history

Cryptic Variation between Species and the Basis of Hybrid Performance

Studies on natural variation in gene expression and its phenotypic effects provide fresh insights into the origins of vigour and sterility in species hybrids

Gene Expression Divergence is Coupled to Evolution of DNA Structure in Coding Regions

Sequence changes in coding region and regulatory region of the gene itself (cis) determine most of gene expression divergence between closely related species. But gene expression divergence between yeast species is not correlated with evolution of primary nucleotide sequence. This indicates that other factors in cis direct gene expression divergence. Here, we studied the contribution of DNA three-dimensional structural evolution as cis to gene expression divergence. We found that the evolution of DNA structure in coding regions and gene expression divergence are correlated in yeast. Similar result was also observed between Drosophila species. DNA structure is associated with the binding of chromatin remodelers and histone modifiers to DNA sequences in coding regions, which influence RNA polymerase II occupancy that controls gene expression level. We also found that genes with similar DNA structures are involved in the same biological process and function. These results reveal the previously unappreciated roles of DNA structure as cis-effects in gene expression