Dissertations / Theses on the topic 'Méthylation d'histone'
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Li, Mengyuan. "Deciphering the roles of Jumonji domain containing proteins in Podospora anserina." Electronic Thesis or Diss., université Paris-Saclay, 2024. http://www.theses.fr/2024UPASL071.
Full textIn eukaryotes, histone proteins associate with DNA in the nucleus to form chromatin. Histone post-translational modifications affect chromatin condensation and gene expression. Histone PTMs are regulated by several enzymes including histone demethylases that often contain a Jumonji C (JmjC) domain. In mammals, JmjC proteins with impaired activity have been shown to cause various developmental defects, metabolic disorders and cancer. Here I present a systematic analysis of the function of the JmjC family of proteins in the model fungus Podospora anserina. The P. anserina genome contains 12 putative lysine demethylase enzymes (Kdm) with two expression patterns, ubiquitous or differentially expressed at different stages of the life cycle. To investigate the function of these Kdms, I generated deletion mutants. My results showed that the Kdms are involved in a wide range of developmental processes in P. anserina. PaKdm4 and PaDmm1 are associated with stress response pathways. Loss of PaKdm1 and PaKdm8-para1 leads to increased longevity of P. anserina. PaKdm5 and PaKdm8-para1 are essential for sexual reproduction. To characterize the consequences of the loss of Kdms on the distribution of H3K4me3, H3K9me3 and H3K36me2/3 histone modifications, I performed chromatin immunoprecipitation on the deleted strains. Loss of PaKdm1 significantly increases H3K4me3 modification, while loss of either PaKdm1 or PaKdm4 results in larger H3K9me3 domains. Performing RNA-seq experiments, I also characterized the transcriptome of the deleted strains. I was able to show that the lack of histone demethylases affects gene expression. Specifically, loss of PaKdm5 has the most significant effect on gene expression, whereas the PaKdm1 deleted strain exhibits limited transcriptional reprogramming. PaKdm8 acts as a positive regulator of gene expression. These findings demonstrate that Kdms epigenetically regulate multiple aspects of the P. anserina life cycle, including lifespan, sexual reproduction and genome defense, and provide new insights into the critical role of histone demethylases in the transcriptional gene regulation
Spadiliero, Barbara. "Epigenetic traits in the parasite Trypanosoma cruzi : DNA and histones modifications linked to its life cycle." Paris 6, 2009. http://www.theses.fr/2009PA066759.
Full textAudonnet, Laure. "Caractérisation fonctionnelle de JMJ24, une déméthylase d’histone de la famille JUMONJI, chez Arabidopsis thaliana." Thesis, Paris 11, 2014. http://www.theses.fr/2014PA112033/document.
Full textNumerous studies over the last decade have reported the characterization of the JUMONJI (JMJ) proteins, showing their critical importance in regulating genes and organism’s development. These proteins are able to demethylate a subset of histone tail residues and were clustered into distinct groups using a phylogenetic analysis based on their catalytic domain conservation. Furthermore, modification of one to three specific residues has been attributed to each JMJ group. Within the KDM3 subfamily, of which target is the H3K9 residue, only one member, IBM1, was first characterized in Arabidopsis. In this report, we showed that the mutation of JMJ24, another member of this subfamily, resulted in an increase of the root length, cotyledon and floral organ size, suggesting that JMJ24 functions is needed at different developmental stage. In addition, the analysis of the tissue-specific expression of JMJ24 indicated that the gene is expressed within the phloem of all organs, correlating with the pleiotropic effect of the gene mutation. Last, our data also suggested that JMJ24 interacts with other JMJ protein like JMJ14 and IBM1, but also with the DCL proteins knowing to be involved in genes and transposable elements regulation
Aranyi, Tamas. "Étude du rôle des facteurs épigénétiques dans la régulation de l'expression du gène de la tyrosine hydroxylase." Paris 6, 2003. http://www.theses.fr/2003PA066007.
Full textGrauffel, Cédric. "Etude in silico de la reconnaissance de la méthylation des lysines sur les queues d'histones." Strasbourg, 2009. http://www.theses.fr/2009STRA6202.
Full textPost-translationnal modifications (PTMs) of histone tails play an important role in cellular processes such as gene expression and regulation. They can act through their specific recognition by proteic domains belonging to chromatin remodeling factors (enzymes, transcription factors, etc). A particularly important modification is the methylation of specific histone tail sites, for which there exist many. Our goal was to get insights into the molecular selectivity of these proteic domains towards a given methylation site by taking advantage of the numerous structural data available in the Protein Data Bank. We studied the interactions between the domains and histone peptides, using molecular dynamics simulations combined with a protocol using the MM/PBSA method. A pre-requisite was to develop and validate force field parameters for the of PTMs. Our results notably revealed that some domains discriminate the methlyation sites by recognizing short linear sequence motifs. It thus appears that chromodomains are specific to an ARKmeS motif, while PHD fingers bind the ARTKme motif of H3. The study carried out on PHD fingers also proves that our overall results can be used as a reference for research of other effectors of histone tails
Touzeau, Amandine. "Identification du chaperon d'histones Spt16 acteur essentiel des réarrangements du génome et méthylation des adénines chez Paramecium tetraurelia." Thesis, Sorbonne Paris Cité, 2018. https://theses.md.univ-paris-diderot.fr/TOUZEAU_Amandine_1_va_20180925.pdf.
Full textIn eukaryotes, chromatin organization is required for the regulation of gene expression and genome stability. Ciliate provide excellent model to study mechanisms involved in maintain of genome integrity. Our study model, the unicellular eukaryote Paramecium tetraurelia, has the particularity to eliminate massively and reproducibly 30% of germinal DNA sequences during the development of the somatic macronucleus after sexual events. Those sequences are eliminated by a multi-step process involving small RNA-directed heterochromatin formation followed by DNA excision by the domesticated transposase Piggy Mac (Pgm) and DNA repair. Molecular mechanisms underlying the specific recognition of those germinal sequences in chromatin context and the precision of the excision, remain elusive. The histone chaperone Spt16, associated to its partner Pob3, is part of the heterodimeric complex FACT (FAcilitates Chromatin Transactions). FACT is implicated in many mechanisms involving DNA metabolism such as transcription, repair, replication or chromatin accessibility. In P. tetraurelia, we identified two homologous proteins to Spt16 and Pob3 expressed only during macronucleus development at the time when genome rearrangements occur. Spt16-1 and Pob3-1 fused to GFP are localized in developing macronuclei. We showed that Spt16-1 is required to obtain a viable sexual progeny. Genome re-sequencing after SPT16-1 inactivation showed that Spt16-1 was required for all DNA elimination events and leads to similar phenotypes and defects to those obtained after PGM inactivation. Spt16-1 acts downstream of small RNA-directed heterochromatin formation and upstream of Pgm. We showed that Spt16-1 was required for the correct localization of Pgm responsible for DNA double strand breaks in developing macronuclei. We proposed a model in which Spt16-1 mediates interaction between chromatin and excision machinery that facilitates access to DNA cleavage sites for the Pgm endonuclease. Adenine DNA methylation, well known in bacteria for its role in restriction modification system, has been described during the last two years in several eukaryotes but in low proportion in the genome. However, its role in eukaryotes remains elusive. Previous analyses by chromatography with radio labelled nucleotides detected around 2,5% of methylated adenine in P. tetraurelia but its precise localization and role have never been analyzed. It has been proposed that this modification could mark with precision the excision sites during genome rearrangements when part of the eliminated sequences carry AT boundaries. The abundance of methylated adenine makes of Paramecium an excellent model to study this modification. Combining immunofluorescence techniques, HPLC-MS and sequencing, we described the presence and the cellular localization during life cycle of 6mA in P. tetraurelia. Those approaches allowed us to show that methylated cytosine are absent in Paramecium. We showed that methylation is mainly found in the somatic genome and transiently in germinal genome. It appears during somatic macronucleus development when genome rearrangements occur. Those preliminary results will allow us to pursue the study of adenine methylation by identifying responsible enzymes and its role in the cell
Zhao, Wei. "Caractérisation moléculaire et fonctionnelle des gènes impliqués dans la mise en place et la lecture de la méthylation d'histones chez l'Arabidopsis thaliana." Thesis, Strasbourg, 2017. http://www.theses.fr/2017STRAJ033/document.
Full textHistone methylation is one of the keys epigenetic marks evolutionarily conserved in eukaryotes. My study focuses on the characterization of factors potentially involved in the deposition and reading of lysine (K) methylation to appreciate its role and underlying mechanisms in the regulation of transcription and plant development, using Arabidopsis thaliana as a model organism. In the first part of my thesis, I report on our study of SET DOMAIN GROUP7 (SDG7), a protein containing the evolutionarily conserved SET domain, which is generally recognized as a signature of K-methyltransferases. We found that SDG7 plays an important role in the regulation of VIN3 induction associated with cold duration measure during vernalization treatment. Intriguingly, levels of several different histone methylations were found unchanged in the sdg7 mutant plants and the recombinant SDG7 protein failed to show a histone-methyltransferase activity in vitro. We thus conclude that SDG7 might methylate a yet unknown non-histone protein to regulate transcription and proper measurement of the duration of cold exposure in the vernalization process. In the second part, I studied interaction between SDG8 and HISTONE MONOUBIQUITINATION1 (HUB1) and HUB2. My results unravel that H3K36me3 and H2Bub1 are deposited largely independently in Arabidopsis, which is in contrast to the dependent crosstalk of these two different epigenetic marks previously reported in yeast. In the last part of my thesis, I report on the identification of the PWWP-domain proteins HUA2/HULK2 as readers of H3K36me3 and demonstrate that sdg8 and hua2 genetically interacts in the regulation of flowering time
Ajjan, Sophie. "Formes atypiques d'empreinte génomique : transitoire, tissu-spécifique et lignée-spécifique." Electronic Thesis or Diss., Paris 6, 2015. http://www.theses.fr/2015PA066251.
Full textGenomic imprinting refers to the functional non-equivalence of the two parental genomes in mammals. Imprinted genes are expressed only from the paternal or maternal allele: this mono-allelic expression is regulated by parent-inherited DNA methylation of specific cis-regulatory regions called ICRs (Imprinting Control Regions). There are currently around 120 imprinted genes known in the mouse genome, which are under the control of 20 characterized ICRs, and are generally conserved in Human. My thesis project aimed at characterizing new maternal ICRs and at analyzing their impact on gene regulation, based on a genome-wide methylation screen conducted in the mouse. I participated to revealing the existence of three forms of genomic imprinting, which reflects variable susceptibility to developmentally-regulated DNA methylation changes: 1) ubiquitous and life-long imprinting, which refers to the 20 canonical ICRs, 2) transient, whose existence is limited to preimplantation development, and 3) tissue-specific. More specifically, I deciphered the histone modification profiles of two new maternal ICR associated with the Cdh15 and the Gpr1/Zdbf2 loci and confirmed that the GPR1/ZDBF2 locus is also subject to transient imprinting in Human. My main achievement concerns the characterization of a candidate ICR associated with the Socs5 gene, which I found to be tissue-specific but also strain-specific, pointing towards a new form of imprinting polymorphism. This ICR has an intragenic position and has the characteristics of an enhancer, hypothesis that I am functionally testing in vivo by a CRISPR/Cas9-mediated deletion. The discovery of these new forms of genomic imprinting provides a better understanding of this phenomenon and its impact on phenotypes
Ajjan, Sophie. "Formes atypiques d'empreinte génomique : transitoire, tissu-spécifique et lignée-spécifique." Thesis, Paris 6, 2015. http://www.theses.fr/2015PA066251/document.
Full textGenomic imprinting refers to the functional non-equivalence of the two parental genomes in mammals. Imprinted genes are expressed only from the paternal or maternal allele: this mono-allelic expression is regulated by parent-inherited DNA methylation of specific cis-regulatory regions called ICRs (Imprinting Control Regions). There are currently around 120 imprinted genes known in the mouse genome, which are under the control of 20 characterized ICRs, and are generally conserved in Human. My thesis project aimed at characterizing new maternal ICRs and at analyzing their impact on gene regulation, based on a genome-wide methylation screen conducted in the mouse. I participated to revealing the existence of three forms of genomic imprinting, which reflects variable susceptibility to developmentally-regulated DNA methylation changes: 1) ubiquitous and life-long imprinting, which refers to the 20 canonical ICRs, 2) transient, whose existence is limited to preimplantation development, and 3) tissue-specific. More specifically, I deciphered the histone modification profiles of two new maternal ICR associated with the Cdh15 and the Gpr1/Zdbf2 loci and confirmed that the GPR1/ZDBF2 locus is also subject to transient imprinting in Human. My main achievement concerns the characterization of a candidate ICR associated with the Socs5 gene, which I found to be tissue-specific but also strain-specific, pointing towards a new form of imprinting polymorphism. This ICR has an intragenic position and has the characteristics of an enhancer, hypothesis that I am functionally testing in vivo by a CRISPR/Cas9-mediated deletion. The discovery of these new forms of genomic imprinting provides a better understanding of this phenomenon and its impact on phenotypes
Battisti, Valentine. "Rôle d'histones methyltransférases spécifiques de H3K9 dans l'équilibre prolifération et différenciation cellulaire." Thesis, Paris 11, 2013. http://www.theses.fr/2013PA11T092/document.
Full textIn eukaryotes, gene expression partly relies on chromatin compaction degree. Chromatin status is controlled by epigenetic marks, such as histones (chromatin structural proteins) posttranslational modifications. As an example, histone H3 lysine 9 (H3K9) methylation on gene promoters is mainly associated with transcriptional repression. H3K9 is methylated by several enzymes called lysine methyltransferases (KMTs). The aim of my thesis project was to understand the role of the H3K9 KMTs, G9a, GLP, Suv39h1 and SETDB1 in regulating the balance between proliferation and terminal differentiation. For this purpose, I used skeletal muscle terminal differentiation as model. Upon muscle terminal differentiation, myoblasts exit, in an irreversible way, from the cell cycle and under go differentiation where cells fusion and form myotubes. During this process, cell cycle genes are permanently silenced and muscle specific genes are activated. Thesis introduction is divided into three chapters. The first chapter focuses on chromatin and post-translational modifications. The second chapter describes H3K9 methylation characteristics and the role of the four KMTs that I studied during my thesis project: G9a,GLP, Suv39h1 and SETDB1. In the third chapter, the skeletal muscle terminal differentiation model is described in details. Results section reports my two major studies outcomes and their discussion. The first concerns the antagonistic roles of G9a and GLP regarding the muscle terminal differentiation and the second focuses on the role of SETDB1 during muscle differentiation. Finally, I conclude this manuscript by a plainer discussion followed by long term perspectives and an appendix presents other research articles, in which I collaborated during my PhD
Marion-Poll, Lucile. "Modifications épigénétiques et transcription dans les deux types de neurones épineux de taille moyenne du striatum." Thesis, Paris 6, 2014. http://www.theses.fr/2014PA066172.
Full textThe striatum is a brain region implicated in physiological functions such as reinforcement learning or movement selection but also in pathologies such as addiction or Parkinson’s disease. It relies on two types of projecting neurons, named “medium spiny neurons” because of their morphology. They are very similar but have a complementary and opposite role. One type expresses the dopamine receptor type 1 (D1R) and the other type expresses the dopamine receptor type 2 (D2R). The aim of this work was to characterize this two neuronal types epigenetically, in basal conditions and after cocaine treatment. We have developed new flow cytometry techniques to be able to distinguish the two cell types. The first method uses transgenic L10-eGFP mice and fresh tissue, the second one goes beyond the limitations of the first one and uses fixed tissue. We have shown that cocaine regulates many post-translational histone modifications, dynamically, and differently between the two populations. Moreover, we have identified more than 100 genes differentially methylated or hydroxymethylated between the two neuronal types. Some of these genes are already known for having a functional role in one of the populations. The comparison between D1R and D2R neurons is a good model to explore the links between DNA methylation, hydroxymethylation and transcription. For example, we have observed a strong association between an increase in DNA methylation and a transcriptional repression, as well as a correlation between DNA methylation and hydroxymethylation
Castillo, Aguilera Omar. "Conception, synthèse et évaluation pharmacologique d’inhibiteurs potentiels de DOT1L impliqués dans la régulation épigénétique du cancer." Thesis, Lille 2, 2017. http://www.theses.fr/2017LIL2S053/document.
Full textCancer is a serious issue of public health as it is one of the main causes of mortality worldwide. Despite the multiple available treatments, it is necessary to develop more efficient and less invasive therapies against cancer. The knowledge of the human genome and epigenome has directed research to new cancer treatment approaches: it is possible to modulate the biological outcome by controlling the access to the genetic information by means of the epigenetic regulation.Epigenetics are the changes happening on the genome without modifying its DNA sequence, leading to a heritable andstable phenotype. In the eukaryotic chromatin, epigenetic regulation implies covalent modifications of DNA and histones. These chemical modifications remodel the chromatin structure leading to an “opened” or “closed” configuration, which is related to the expression or repression of genes. The epigenetic landscape is altered in cancers; for example, abnormal methylation leads to the silencing of certain genes (such as tumor suppressor genes), or to the over-expression of oncogenes. Unlike genetic alterations that are irreversible, epigenetic aberrations are reversible. Thus, molecules that can reestablish the epigenetic balance represent potent therapeutic tools for cancer treatment.Methylation and acetylation are the most studied epigenetic modifications. DNA methylation is carried out by the DNAmethyltransferases (DNMTs) and histone methylation by the histone methyltransferases (HMTs).This PhD project was focused on the histone methyltransferase DOT1L (DOT1 like, disruptor of telomeric silencing), responsible of methylation of residue Lys79 of histone 3 (H3K79), which leads to the transcription of some oncogenes. Recent studies have shown that DOT1L is implicated in MLL-rearranged leukemia (MLL-r, Myeloid-Lymphoid Leukemia) thus it is a potent target in cancer. As DOT1L and DNMTs share the same cofactor, S-adenosyl-L-methionine (SAM), DNMT and DOT1L inhibitors can present a common inhibition mechanism by competing with SAM.We present herein the in silico – based design, and the multi-step synthesis of some series of molecules containing 3 main moieties: a) an aminopyrimidine motif and b) a benzimidazole or phenylurea motif, linked by c) a phenyl or heterocycle motif. DOT1L activity was determined for the different compounds synthesized and structure-activity relationships (SAR) were established. The activity on DNMT and other HMTs was determined as well, in other to study the DOT1L specificity of our compounds.Different scaffolds were identified to obtain DOT1L-selective or DOT1L/DNMT dual inhibitors. These molecules are interesting therapeutic tools for cancer treatment
Oréal, Vincent. "Rôles distincts des différentes formes de méthylation de H3K4 dans deux mécanismes de répression transcriptionnelle et mise en évidence d'une nouvelle voie de surveillance moléculaire liée à l'excès d'histones libres." Thesis, Aix-Marseille 2, 2010. http://www.theses.fr/2010AIX22051/document.
Full textRelationships between histones, components of nucleosomes, and the transcription process of coding genes are both multiple and extremely complex. During my Thesis, I looked at twoof these relationships. First, we performed a study in collaboration with the Franck Holstedge and Catherine Dargemont labs. This work has allowed us to clearly define the effect of various methylation forms of the lysine 4 of the histone 3 on gene transcription. In this study we have shownthat H3K4 methylation influences the transcription of only a very limited number of genes. For these genes, a non conventional distribution profile of H3K4 methylation forms has been identified by the presence of an unusual enrichment in di- and trimethylated H3K4 in the 3’ of these genes. The principal effect of this mark is to promote transcriptional repression by at least two distinct mechanisms. The atypical enrichment of H3K4 trimethylation negatively influences gene expressionvia the production of non coding antisense RNA. For the repressive effect associated with dimethylH3K4, the quantity of antisense RNA as well as its production are not involved. We propose severalh ypotheses that link our results to the data known on this subject. In a second study performed incollaboration with the Sebastian Chavez and Akash Gunjan labs, we concentrated on the FACT complex that is involved in the assembly and disassembly of nucleosomes as RNA polymerase IImoves past. Previously, a growth defect in thermosensitive mutants of the FACT complex had been observed. In our study, we show that FACT deterioration leads to the eviction of histones that arenormally incorporated into chromatin during transcription. The accumulation of these free histones,which have a high toxic potential, induces the specific repression of CLN3 which encodes for the firstcyclin of G1 phase. For the first time, we show in this study the existence of a cell cycle molecular surveillance mechanism that is induced by an excess of free histones
Zhang, Xue. "Implications fonctionnelles de deux histone méthyltransferases dans les réponses aux stress et la régulation de la floraison chez Arabidopsis thaliana." Thesis, Strasbourg, 2019. http://www.theses.fr/2019STRAJ074.
Full textHistone methylation catalyzed by histone methyltransferase is essential in transcriptional regulation of gene expression. Histone methyltransferases are known to play crucial roles in multiple cellular processes in plants. My PhD work investigated the biological function of two histone methyltransferases in controlling plant responses to various environmental stimuli in Arabidopsis thaliana. In the first part, my results demonstrated that the H3K36-methyltransferase SDG8 transcriptionally regulates NPR1, a central player in salicylic acid-mediated immunity and co-acts with the RNAPII to enable the efficient transcriptional induction of several defense genes upon stimulation. In the second part, my work unraveled that SDG26, another ortholog of the animal H3K36-methyltransferase, plays an important role in plant response to abiotic stresses. By focusing on cold stress, SDG26 was shown to regulate the cold stress response by directly activating the transcription of SOC1 and CBF genes through binding their chromatin and depositing H3K36me3. Interestingly, SDG26 mastered the accumulation of ABA by regulating the expression of ABA homeostasis-related genes, suggesting an involvement of ABA pathway in the cold response. In the last part, using a genetic approach my work established SDG26 as an autonomous flowering pathway component. Accordingly, SDG26 was found in a multiple-protein complex comprising the histone demethylase FLD, the homeobox-domain transcription factor LD, as well as a putative COMPASS component APRF1. This multiple-protein complex was found in controlling the repression of the major flowering repressor FLC as well as the activation of the flowering activator SOC1 to precisely regulate the floral transition
Broucqsault, Natacha. "Régulation épigénétique du locus subtélomérique 4q35 et contribution du macrosatellite D4Z4 dans la dystrophie facio-scapulo-humérale." Thesis, Aix-Marseille, 2013. http://www.theses.fr/2013AIXM5099.
Full textHird more frequently myopathy, the facioscapulohumeral dystrophy is an enigmatic disease. It is associated with the macrosatellite D4Z4 contraction since 90’s but the origin of the modifications observable in patients remains unclear. That’s why, we have focused our work in different aspects of this disease. First of all, we have studied expression of 4q35 and muscular genes in fetuses and adults carrying a contraction of the D4Z4 macrosatellite and shown that the molecular deregulations can be observed since the fetal stage. So, we have validated this model as an interesting model to study the early FSHD pathogenesis. Secondly, we have been interested in epigenetic regulation of two genes located in the 4q35 region and have observed a modulation of their expression in a global epigenetic modulation contest. Those deregulations depend on the number of D4Z4 repeats. Finally, we have analyzed the region regulation by telomere position effect and have noted only some genes are deregulated by this epigenetic mechanism. With those results, we have progressed in the FSHD pathogenesis knowledge and we have validated a new model to study this pathology
Filleton, Fabien. "Cartographie et analyse de variations épigénomiques naturelles chez la levure Saccharomyces cerevisiae." Thesis, Lyon, École normale supérieure, 2015. http://www.theses.fr/2015ENSL1044.
Full textEpigenome is defined as the entire chromatin information other than the DNA sequence. Within a given species and for a given cell type, each indivual has specific epigenomic characteristics. Epigenomic differences between individuals (refered to as 'epi-polymorphisms') remain poorly characterized, although cases were reported where they could be linked to phenotypic differences. In my thesis, I used the model organism S. cerevisiae to identify histone modification epi-polymorphisms and study their biological impact. I profiled the epigenome of five different histone modifications (3 acetylations and 2 methylations) in three natural yeast strains. By ChIP-seq methods and software developments, I compared these strains at single-nucleosome resolution and discovered novel characteristics of these epi-polymorphisms which are described in this manuscript.Furthermore, I constructed a research framework to investigate the link between epi-polimorphisms and response to environmental cues. For this, I built a set of mutant strains derived from natural strains but where some epi-polymorphisms can no longer be maintained. I analyzed by RNA-seq the transcriptomes of some of these mutant strains before and after an environmental shift. Unfortunately, the quality of this initial data produced was not sufficient to link epi-polymorphisms to differntial responses, but the strain resources remain available for further investigations. Finally, I studied the evolutionary dynamics of epi-polymorphisms in the presence or absence of selection pressure. To do so, I followed the evolution of H3K14ac for 1.000 generations under two conditions of yeast experimental evolution ( selective or neutral). Marked differences were observed between the two regimes, revealing unexpected consequences of the presence of selection. Further mechanistic studies will be needed to elucidate the full properties of these differences