@misc{20833,
  abstract     = {Sex-chromosome systems are highly variable across animals, but how they transition from one to another is not well understood. Diptera have undergone multiple sex-chromosome turnovers and expansions while maintaining their general chromosomal content, which makes them an ideal clade to study such transitions. We analysed more than 100 dipteran whole-genome assemblies and identified 4 new lineages that underwent sex-chromosome turnover (in addition to the 5 previously reported). We find the majority of turnovers happened in the group Schizophora, which tend to have fewer genes on the F element (the chromosome homologous to the ancestral insect X chromosome) than lower dipterans, a factor previously hypothesized to facilitate turnover. Most derived X chromosomes have higher GC content than autosomes, consistent with a high prevalence of male-achiasmy in Diptera. In addition, an excess of gene movement out of the X is detected for most of these new X chromosomes, and many of these moved genes have high testis expression in Drosophila, suggesting that out-of-X gene movement contributes to the long-term demasculinization of X chromosomes.},
  author       = {Layana Franco, Lorena Alexandra and Toups, Melissa A and Vicoso, Beatriz},
  keywords     = {Schizophora, sex chromosomes, sex-chromosome turnover, Diptera, genomic features, out-of-X movement.},
  publisher    = {Institute of Science and Technology Austria},
  title        = {{Research Data for 'Causes and consequences of sex-chromosome turnovers in Diptera'}},
  doi          = {10.15479/AT-ISTA-20833},
  year         = {2026},
}

@misc{21116,
  abstract     = {Sex-chromosome systems are highly variable across animals, but how they transition from one to another is not well understood. Diptera have undergone multiple sex-chromosome turnovers and expansions while maintaining their general chromosomal content, which makes them an ideal clade to study such transitions. We analyzed more than 100 dipteran whole-genome assemblies and identified 4 new lineages that underwent sex-chromosome turnover (in addition to the 5 previously reported). We find that the majority of turnovers happened in the group Schizophora, which tend to have fewer genes on Muller element F (the chromosome homologous to the ancestral insect X chromosome) than lower dipterans, a factor previously hypothesized to facilitate turnover. Most derived X chromosomes have higher GC content than autosomes, consistent with a high prevalence of male achiasmy in Diptera. In addition, an excess of gene movement out of the X is detected for most of these new X chromosomes, and many of these moved genes have high testis expression in Drosophila, suggesting that out-of-X gene movement contributes to the long-term demasculinization of X chromosomes.},
  author       = {Layana Franco, Lorena Alexandra and Toups, Melissa A and Vicoso, Beatriz},
  keywords     = {Schizophora, sex chromosomes, sex-chromosome turnover, Diptera, genomic features, out-of-X movement.},
  publisher    = {Institute of Science and Technology Austria},
  title        = {{Research Data for "Causes and consequences of sex-chromosome turnovers in Diptera"}},
  doi          = {10.15479/AT-ISTA-21116},
  year         = {2026},
}

@article{21409,
  abstract     = {Meiotic drivers are selfish genetic elements that gain transmission advantages by distorting equal, Mendelian segregation. For decades, biologists have considered meiotic drivers as interesting, albeit esoteric, case studies. It is now clear, however, that meiotic drive is more common and phylogenetically widespread than previously supposed. Indeed, intensive study of a few well-known cases has begun to reveal the evolutionary genomic consequences of meiotic drive. We argue here that many features of genome evolution, content, and organization that are seemingly inexplicable by organismal adaptation or nearly neutral processes are instead best accounted for by recurrent histories of meiotic drive. We review how meiotic drive can affect the evolution of sequences, gene copy numbers, genes with functions in meiosis and gametogenesis, signatures of “selection,” chromosome rearrangements, and karyotype evolution. We also explore the interactions of meiotic drive elements with other classes of selfish genetic elements, including satellite DNAs, transposable elements, and with the endogenous host genes involved in drive suppression. Finally, we argue that some aspects of drive-mediated genome evolution are now sufficiently well established that we might reverse the direction of discovery—rather than ask how drive affects genome evolution, we can use genome data to discover new putative drive elements.},
  author       = {Presgraves, Daven C. and Dawe, R. Kelly and Dyer, Kelly A. and Fishman, Lila and Bhide, Soumitra A. and Bradshaw, Sasha L. and Brady, Meghan J. and Burga, Alejandro and Courret, Cécile and Fagen, Brandon L. and Machado Ferretti, Ana Beatriz Stein and Kelemen, Réka K and Kitano, Jun and Liu, Yiran and Martí, Emiliano and Erlenbach, Theresa and Reinhardt, Josephine A. and Ross, Laura and Runge, Jan Niklas and Swanepoel, Callie M. and Vicoso, Beatriz and Vogan, Aaron A. and Lindholm, Anna K. and Larracuente, Amanda M. and Unckless, Robert L.},
  issn         = {1537-1719},
  journal      = {Molecular Biology and Evolution},
  number       = {2},
  publisher    = {Oxford University Press},
  title        = {{The evolutionary genomics of meiotic drive}},
  doi          = {10.1093/molbev/msag020},
  volume       = {43},
  year         = {2026},
}

@article{21161,
  abstract     = {In many species, sex-biased expression is widespread and thought to contribute to sexual dimorphism. While bulk RNA-sequencing has been instrumental in identifying strongly sex-biased genes, it lacks resolution to assess variation across cell-types and tissue compartments. Using single-nucleus expression data from the Fly Cell Atlas, we investigate sex differences in adult Drosophila melanogaster. We find that differences in cell-type composition between the sexes are not a major source of sex-bias, as for the vast majority of genes, the degree of sex-bias is similar regardless of whether sex differences in cell-type composition are controlled for or not. Our analysis confirms a deficit of X-linked male-biased genes in the body’s somatic tissues that is widespread across cell-types. We also find the excess of X-linked female-biased genes to be associated with nervous system cells in the head but with epithelial cells in the body’s somatic tissues, showing that single-nucleus data crucially resolves sex-bias at the cell-type level. We investigate dosage compensation (DC) across 15 tissues and 17 cell-types. We observe that it varies throughout the body. Surprisingly, we observe a lack of DC in a cluster of main cells within the male accessory glands. This result highlights the importance of understanding context-dependent DC.},
  author       = {De Castro Barbosa Rodrigues Barata, Carolina and Vicoso, Beatriz},
  issn         = {1471-2954},
  journal      = {Proceedings of the Royal Society B Biological Sciences},
  number       = {2063},
  publisher    = {Royal Society of London},
  title        = {{Single-nucleus resolution of sex-biased expression and dosage compensation in Drosophila melanogaster}},
  doi          = {10.1098/rspb.2025.2471},
  volume       = {293},
  year         = {2026},
}

@article{20655,
  abstract     = {Traits that affect organismal fitness are often highly genetically variable. This genetic variation is vital for populations to adapt to their environments, but it is also surprising given that nature – after all – ‘selects’ the best genotypes at the expense of those that fall short. Explaining the extensive genetic variation of fitness‐related traits is thus a longstanding puzzle in evolutionary biology, with cascading implications for ecology, conservation, and human health. Balancing selection – an umbrella term for scenarios in which natural selection maintains genetic variation – is a century‐old explanation to resolve this puzzle that has gained recent momentum from genome‐scale methods for detecting it. Yet evaluating whether balancing selection can, in fact, resolve the puzzle is challenging, given the logistical constraints of distinguishing balancing selection from alternative hypotheses and the daunting collection of theoretical models that formally underpin this debate. Here, we track the development of balancing selection theory over the last century and provide an accessible review of this rich collection of models. We first outline the range of biological scenarios that can generate balancing selection. We then examine how fundamental features of genetic systems – non‐random mating between individuals, ploidy levels, genetic drift, linkage, and genetic architectures of traits – have been progressively incorporated into the theory. We end by linking these theoretical predictions to ongoing empirical efforts to understand the evolutionary processes that explain genetic variation.},
  author       = {Ruzicka, Filip and Zwoinska, Martyna K. and Goedert, Debora and Kokko, Hanna and Li Richter, Xiang‐Yi and Moodie, Iain R. and Nilén, Sofie and Olito, Colin and Svensson, Erik I. and Czuppon, Peter and Connallon, Tim},
  issn         = {1469-185X},
  journal      = {Biological Reviews},
  keywords     = {evolutionary theory, population genetics, balancing selection, heterozygote advantage, trade-offs, negative frequency-dependent selection, fitness variation, mathematical modelling},
  number       = {2},
  publisher    = {Wiley},
  title        = {{A century of theories of balancing selection}},
  doi          = {10.1111/brv.70103},
  volume       = {101},
  year         = {2026},
}

@article{21486,
  abstract     = {Sex-chromosome systems are highly variable across animals, but how they transition from one to another is not well understood. Diptera have undergone multiple sex-chromosome turnovers and expansions while maintaining their general chromosomal content, which makes them an ideal clade to study such transitions. We analyzed more than 100 dipteran whole-genome assemblies and identified 4 new lineages that underwent sex-chromosome turnover (in addition to the 5 previously reported). We find that the majority of turnovers happened in the group Schizophora, which tend to have fewer genes on Muller element F (the chromosome homologous to the ancestral insect X chromosome) than lower dipterans, a factor previously hypothesized to facilitate turnover. Most derived X chromosomes have higher GC content than autosomes, consistent with a high prevalence of male achiasmy in Diptera. In addition, an excess of gene movement out of the X is detected for most of these new X chromosomes, and many of these moved genes have high testis expression in Drosophila, suggesting that out-of-X gene movement contributes to the long-term demasculinization of X chromosomes.},
  author       = {Layana Franco, Lorena Alexandra and Toups, Melissa A and Vicoso, Beatriz},
  issn         = {2056-3744},
  journal      = {Evolution Letters},
  number       = {3},
  publisher    = {Oxford University Press},
  title        = {{Causes and consequences of sex-chromosome turnovers in Diptera}},
  doi          = {10.1093/evlett/qrag003},
  volume       = {10},
  year         = {2026},
}

@article{21900,
  abstract     = {Individually silencing 125 fruit fly genes reveals opposing fitness effects of mutations between females and males, as well as between germline and somatic tissues.},
  author       = {Ruzicka, Filip},
  issn         = {2397-334X},
  journal      = {Nature Ecology & Evolution},
  pages        = {1035--1036},
  publisher    = {Springer Nature},
  title        = {{Reverse genetics of sexual antagonism}},
  doi          = {10.1038/s41559-026-03036-y},
  volume       = {10},
  year         = {2026},
}

@article{20009,
  abstract     = {The suppression of recombination between young X and Y chromosomes is a crucial step in their evolution, but why it occurs is not known. The detailed characterization of the polymorphic sex chromosomes of the fourspine stickleback by Liu et al. promises to shed new light on this longstanding question.},
  author       = {Vicoso, Beatriz},
  issn         = {0169-5347},
  journal      = {Trends in Ecology and Evolution},
  number       = {8},
  pages        = {728--730},
  publisher    = {Elsevier},
  title        = {{Sex chromosome evolution in action in fourspine sticklebacks}},
  doi          = {10.1016/j.tree.2025.06.010},
  volume       = {40},
  year         = {2025},
}

@article{20044,
  abstract     = {Genetic trade-offs—which occur when variants that are beneficial in some contexts of natural selection are harmful in others—can influence a wide range of evolutionary phenomena, from the maintenance of genetic variation to the evolution of aging and sex differences. An extensive body of evolutionary theory has focused on the consequences of such trade-offs, and recent analyses of Fisher’s geometric model have further quantified the expected proportion of new mutations that exhibit trade-offs. However, the theory remains silent regarding the prevalence of trade-offs among the variants that contribute to adaptation. Here, we extend Fisher’s geometric model to predict the prevalence of trade-offs among the adaptive mutations that become established or fixed in a population. We consider trade-offs between sexes, habitats, fitness components, and temporally fluctuating environments. In all 4 scenarios, trade-off alleles are consistently under-represented among established relative to new beneficial mutations—an effect that arises from the greater susceptibility of trade-off alleles to genetic drift. Adaptation during a population size decline exacerbates this deficit of trade-offs among established mutations, whereas population expansions dampen it. Consequently, threatened populations should primarily adapt using unconditionally beneficial alleles, while invasive populations are more prone to adaptation using variants that exhibit trade-offs.},
  author       = {Connallon, Tim and Czuppon, Peter and Olito, Colin and Goedert, Debora and Kokko, Hanna and Nava-Bolaños, Angela and Nilén, Sofie and Svensson, Erik I and Zwoinska, Martyna and Dutoit, Ludovic and Ruzicka, Filip},
  issn         = {1558-5646},
  journal      = {Evolution},
  number       = {7},
  pages        = {1243--1255},
  publisher    = {Oxford University Press},
  title        = {{Predicting the prevalence of genetic trade-offs among adaptive substitutions}},
  doi          = {10.1093/evolut/qpaf061},
  volume       = {79},
  year         = {2025},
}

@article{20182,
  abstract     = {Sex chromosomes have evolved many times throughout the tree of life, and understanding what has shaped their unusual morphological, sequence, and regulatory features has been a long-standing goal. Most early insights into insect sex chromosome biology came from a few model species, such as the fruit fly Drosophila melanogaster, which limited broad-scale evolutionary inferences. More recently, extensive comparative genomics studies have uncovered several unexpected patterns, which we highlight in this review. First, we describe the conservation of the ancestral X chromosome over 450 million years but also its recurrent turnover (i.e. its reversal to an autosome when a new X chromosome arose) in at least one order. We then summarize classical and more recent findings on how insects modulate the expression of X-linked genes following the degradation of the Y chromosome and how the diverse mechanisms of dosage compensation identified may elucidate important principles of sex chromosome regulatory evolution.},
  author       = {Toups, Melissa A and Vicoso, Beatriz},
  issn         = {2214-5753},
  journal      = {Current Opinion in Insect Science},
  publisher    = {Elsevier},
  title        = {{Insect sex chromosome evolution: Conservation, turnover, and mechanisms of dosage compensation}},
  doi          = {10.1016/j.cois.2025.101411},
  volume       = {72},
  year         = {2025},
}

@article{20223,
  abstract     = {The first influential hypothesis for sex chromosome evolution was proposed in 1914 by H. J. Muller, who argued that once recombination was suppressed between the X and Y chromosomes, Y-linked genes become “sheltered” from selection, leading to accumulation of recessive loss-of-function (LOF) mutations and decay of Y-linked genes. The hypothesis fell out of favor in the 1970s because early mathematical models failed to support it and data on the dominance of lethal mutations were viewed as incompatible with the hypothesis. We reevaluate the main arguments against Muller's hypothesis and find that they do not conclusively exclude a role for sheltering in sex chromosome evolution. By relaxing restrictive assumptions of earlier models, we show that sheltering promotes fixation of LOF mutations with sexually dimorphic fitness effects, resulting in decay of X-linked genes that are exclusively expressed by males and Y-linked genes that are primarily, though not necessarily exclusively, expressed by females. We further show that drift and other processes contributing to Y degeneration (i.e. selective interference and regulatory evolution) expand conditions of Y-linked gene loss by sheltering. The actual contribution of sheltering to sex chromosome evolution hinges upon the distribution of dominance and sex-specific fitness effects of LOF mutations, which we discuss.},
  author       = {Mrnjavac, Andrea and Vicoso, Beatriz and Connallon, Tim},
  issn         = {1537-1719},
  journal      = {Molecular Biology and Evolution},
  number       = {8},
  publisher    = {Oxford University Press},
  title        = {{An extension of Muller's sheltering hypothesis for the evolution of sex chromosome gene content}},
  doi          = {10.1093/molbev/msaf177},
  volume       = {42},
  year         = {2025},
}

@article{20330,
  abstract     = {The evolution of sexual dimorphism (the difference in average trait values between females and males, SD), is often thought to be constrained by shared genetic architecture between the sexes. Indeed, it is commonly expected that SD should negatively correlate with the intersex correlation (the genetic correlation between effects of segregating variants in females and males, r fm), either because (1) traits with ancestrally low r fm are less constrained in their ability to respond to sex-specific selection and thus evolve to be more dimorphic, or because (2) sex-specific selection, driving sexual dimorphism evolution, also acts to reduce r fm. Despite the intuitive appeal and prominence of these ideas, their generality and the conditions in which they hold remain unclear. Here, we develop models incorporating sex-specific stabilizing selection, mutation and genetic drift to examine the relationship between r fm and SD. We show that the two commonly-discussed mechanisms with the potential to generate a negative correlation between SD and r fm could just as easily generate a positive association, since the standard line of reasoning hinges on a hidden assumption that sex-specific adaptation more frequently favors increased dimorphism than reduced dimorphism. Our results provide, to our knowledge, the first mechanistic framework for understanding the conditions under which a correlation between r fm and SD may arise and offer a compelling explanation for inconsistent empirical evidence. We also make the intriguing observation that—even when selection between the two sexes is identical—drift generates nonzero SD. We quantify this effect and discuss its significance.},
  author       = {Puixeu Sala, Gemma and Hayward, Laura},
  issn         = {1943-2631},
  journal      = {Genetics},
  number       = {3},
  publisher    = {Oxford University Press},
  title        = {{The relationship between sexual dimorphism and intersex correlation: Do models support intuition?}},
  doi          = {10.1093/genetics/iyaf175},
  volume       = {231},
  year         = {2025},
}

@article{20404,
  abstract     = {Collagens are fundamental components of extracellular matrices, requiring precise intracellular post-translational modifications for proper function. Among the modifications, prolyl 4-hydroxylation is critical to stabilise the collagen triple helix. In humans, this reaction is mediated by collagen prolyl 4-hydroxylases (P4Hs). While humans possess three genes encoding these enzymes (P4H⍺s), Drosophila melanogaster harbour at least 26 candidates for collagen P4H⍺s despite its simple genome, and it is poorly understood which of them are actually working on collagen in the fly. In this study, we addressed this question by carrying out thorough bioinformatic and biochemical analyses. We demonstrate that among the 26 potential collagen P4H⍺s, PH4⍺EFB shares the highest homology with vertebrate collagen P4H⍺s. Furthermore, while collagen P4Hs and their substrates must exist in the same cells, our transcriptomic analyses at the tissue and single cell levels showed a global co-expression of PH4⍺EFB but not the other P4H⍺-related genes with the collagen IV genes. Moreover, expression of PH4⍺EFB during embryogenesis was found to precede that of collagen IV, presumably enabling efficient collagen modification by PH4⍺EFB. Finally, biochemical assays confirm that PH4⍺EFB binds collagen, supporting its direct role in collagen IV modification. Collectively, we identify PH4⍺EFB as the primary and potentially constitutive prolyl 4-hydroxylase responsible for collagen IV biosynthesis in Drosophila. Our findings highlight the remarkably simple nature of Drosophila collagen IV biosynthesis, which may serve as a blueprint for defining the minimal requirements for collagen engineering.},
  author       = {Ishikawa, Yoshihiro and Toups, Melissa A and Elkrewi, Marwan N and Zajac, Allison L. and Horne-Badovinac, Sally and Matsubayashi, Yutaka},
  issn         = {1569-1802},
  journal      = {Matrix Biology},
  number       = {11},
  pages        = {101--113},
  publisher    = {Springer Nature},
  title        = {{Evidence for the major role of PH4⍺EFB in the prolyl 4-hydroxylation of Drosophila collagen IV}},
  doi          = {10.1016/j.matbio.2025.09.002},
  volume       = {141},
  year         = {2025},
}

@misc{20780,
  abstract     = {Sex-chromosome systems are highly variable across animals, but how they transition from one to another is not well understood. Diptera have undergone multiple sex-chromosome turnovers and expansions while maintaining their general chromosomal content, which makes them an ideal clade to study such transitions. We analysed more than 100 dipteran whole-genome assemblies and identified 4 new lineages that underwent sex-chromosome turnover (in addition to the 5 previously reported). We find the majority of turnovers happened in the group Schizophora, which tend to have fewer genes on the F element (the chromosome homologous to the ancestral insect X chromosome) than lower dipterans, a factor previously hypothesized to facilitate turnover. Most derived X chromosomes have higher GC content than autosomes, consistent with a high prevalence of male-achiasmy in Diptera. In addition, an excess of gene movement out of the X is detected for most of these new X chromosomes, and many of these moved genes have high testis expression in Drosophila, suggesting that out-of-X gene movement contributes to the long-term demasculinization of X chromosomes.},
  author       = {Layana Franco, Lorena Alexandra and Toups, Melissa A and Vicoso, Beatriz},
  keywords     = {Schizophora, sex chromosomes, sex-chromosome turnover, Diptera, genomic features, out-of-X movement.},
  publisher    = {Institute of Science and Technology Austria},
  title        = {{Causes and consequences of sex-chromosome turnovers in Diptera}},
  doi          = {10.15479/AT-ISTA-20780},
  year         = {2025},
}

@article{20796,
  abstract     = {Rapid prophase chromosome movements ensure faithful alignment of the parental homologous chromosomes and successful synapsis formation during meiosis. These movements are driven by cytoplasmic forces transmitted to the nuclear periphery, where chromosome ends are attached through transmembrane proteins. During many developmental stages a specific genome architecture with chromatin nuclear periphery contacts mediates specific gene expression. Whether chromatin is removed from the nuclear periphery as a consequence of chromosome motions or by a specific mechanism is not fully understood. Here, we identify a mechanism to remove chromatin from the nuclear periphery through vaccinia related kinase (VRK-1)–dependent phosphorylation of Barrier to Autointegration Factor 1 (BAF-1) in Caenorhabditis elegans early prophase of meiosis. Interfering with chromatin removal delays chromosome pairing, impairs synapsis, produces oocytes with abnormal chromosomes and elevated apoptosis. Long read sequencing reveals deletions and duplications in offspring lacking VRK-1 underscoring the importance of the BAF-1–VRK-1 module in preserving genome stability in gametes during rapid chromosome movements.},
  author       = {Paouneskou, Dimitra and Baudrimont, Antoine and Kelemen, Réka K and Elkrewi, Marwan N and Graf, Angela and Moukbel Ali Aldawla, Shehab and Kölbl, Claudia and Tiemann-Boege, Irene and Vicoso, Beatriz and Jantsch, Verena},
  issn         = {2041-1723},
  journal      = {Nature Communications},
  publisher    = {Springer Nature},
  title        = {{BAF-1–VRK-1 mediated release of meiotic chromosomes from the nuclear periphery is important for genome integrity}},
  doi          = {10.1038/s41467-025-65420-9},
  volume       = {16},
  year         = {2025},
}

@article{19370,
  abstract     = {Sex-linked and autosomal loci experience different selective pressures and evolutionary dynamics. X (or Z) chromosomes are often hemizygous in males (or females), as Y (or W) chromosomes often degenerate. Such hemizygous regions can be under greater efficacy of selection, as recessive mutations are immediately exposed to selection in the heterogametic sex leading to faster adaptation and faster divergence on the X chromosome (the so-called Faster-X or Faster-Z effect). However, in young nonrecombining regions, Y/W chromosomes often have many functional genes, and many X/Z-linked loci are therefore diploid. The sheltering of recessive mutations on the X/Z by the Y/W homolog is expected to drive slower adaptation for diploid X/Z loci, i.e. a reduction in the efficacy of selection. While the Faster-X effect has been studied extensively, much less is known empirically about the evolutionary dynamics of diploid X or Z chromosomes. Here, we took advantage of published population genomic data in the female-heterogametic human parasite Schistosoma japonicum to characterize the gene content and diversity levels of the diploid and hemizygous regions of the Z chromosome. We used different metrics of selective pressures acting on genes to test for differences in the efficacy of selection in hemizygous and diploid Z regions, relative to autosomes. We found consistent patterns suggesting reduced Ne, and reduced efficacy of purifying selection, on both hemizygous and diploid Z regions. Moreover, relaxed selection was particularly pronounced for female-biased genes on the diploid Z, as predicted by recent theoretical work.},
  author       = {Mrnjavac, Andrea and Vicoso, Beatriz},
  issn         = {1759-6653},
  journal      = {Genome Biology and Evolution},
  number       = {2},
  publisher    = {Oxford University Press},
  title        = {{Reduced efficacy of selection on a young Z chromosome region of schistosoma japonicum}},
  doi          = {10.1093/gbe/evaf021},
  volume       = {17},
  year         = {2025},
}

@phdthesis{19386,
  abstract     = {Crustaceans are a large group of arthropods with a great diversity of species and
different types of sex determination systems and reproductive modes (Subramoniam, 2017).
This makes them a great model for exploring the evolution of sex chromosomes and sexual
dimorphism and investigating the evolutionary mechanisms driving and maintaining the
diversity of reproductive systems. Within this taxon, Brine shrimp of the genus Artemia, a
branchiopod crustacean, are well suited for such explorations, as they have both highly
dimorphic traits and closely related sexual and asexual species. Although brine shrimp are
known to have ZW sex chromosomes (Bowen, 1963; Parraguez et al., 2009), the sex
chromosomes are still not well characterized at the genomic level, the sex-determination gene
is unknown, and it is still unclear whether the same sex chromosomes as shared by the
different species.
The first part of this thesis was to characterize the Z and W chromosomes in Artemia
using an array of methods, from generating multiple chromosome and contig level genome
assemblies to identifying W-linked scaffolds and transcripts in multiple species using k-mer
based approaches.
The second part tackles the conservation of the cell type specific regulatory pathways
in the female reproductive system between Artemia and Drosophila, and the expression of the
Z-specific region throughout meiosis using single-nucleus RNA-seq data. Our results show
that germline cells lack dosage compensation, with a subset of cells showing evidence of
extreme repression of the Z chromosome.
With multiple sexual species and several asexual lineages of parthenogenetic females
that produce rare males at low frequencies, Brine shrimp present the perfect opportunity to
explore the transition to asexuality and shed light on the prerequisites and repercussions of
the form of modified meiosis maintaining the asexual lineages. The last chapter is an
investigation of the molecular pathways involved in asexual reproduction in Artemia using
newly generated single nucleus RNAseq and WGS data and previously published data. },
  author       = {Elkrewi, Marwan N},
  isbn         = {9783990780534},
  issn         = {2663-337X},
  pages        = {170},
  publisher    = {Institute of Science and Technology Austria},
  title        = {{Evolution of sex chromosomes, sex determination and asexuality in Artemia brine shrimp}},
  doi          = {10.15479/AT-ISTA-19386},
  year         = {2025},
}

@phdthesis{20449,
  abstract     = {Males and females of many  species differ in morphology, physiology, and behavior. In taxa
with genetic sex determination, sexual differentiation arises largely from sex-biased gene
expression, which varies across tissues, developmental stages, and lineages. Increasing
evidence highlights chromatin configuration, which can exist in open or closed states, and can
be shaped by sex-determination path ways, as a key regulatory layer of this dimorphism.
Degeneration of the Y or W chromosome further contributes to sex -specific differences by
altering gene copy numbers relative to autosomes in heterogametic sex. To mitigate these
imbalances, many eukaryotes have independently evolved dosage compensation mechanisms,
often mediated through chromatin -level regulation. In this thesis, we investigate the
evolutionary dynamics of sex chromosome differentiation in two species, Artemia franciscana
and Cameraria  ohridella , with a particular focus on the extent of dosage compensation
following gene loss in the heterogametic sex and the potential chromatin-based mechanisms
underlying this process. We further characterize sex -biased gene expression and its regulation
through histone modifications. Our analyses also reveal that the A. franciscana genome is
highly repetitive, with many genes containing intronic transposable elements. We find that
enrichment of histonemo difications associated with constitutive heterochromatin, positively
correlates with variation in gene expression levels. Collectively, these findings underscore role
of chromatin regulation in shaping the evolution of sex chromosomes and sexual
differentiation. },
  author       = {Bett, Vincent K},
  issn         = {2663-337X},
  pages        = {114},
  publisher    = {Institute of Science and Technology Austria},
  title        = {{Evolution and regulation of the Z chromosome}},
  doi          = {10.15479/AT-ISTA-20449},
  year         = {2025},
}

@article{19735,
  abstract     = {The males and females of the brine shrimp Artemia franciscana are highly dimorphic, and this dimorphism is associated with substantial sex-biased gene expression in heads and gonads. How these sex-specific patterns of expression are regulated at the molecular level is unknown. A. franciscana also has differentiated ZW sex chromosomes, with complete dosage compensation, but the molecular mechanism through which compensation is achieved is unknown. Here, we conducted CUT&TAG assays targeting 7 post-translational histone modifications (H3K27me3, H3K9me2, H3K9me3, H3K36me3, H3K27ac, H3K4me3, and H4K16ac) in heads and gonads of A. franciscana, allowing us to divide the genome into 12 chromatin states. We further defined functional chromatin signatures for all genes, which were correlated with transcript level abundances. Differences in the occupancy of the profiled epigenetic marks between sexes were associated with differential gene expression between males and females. Finally, we found a significant enrichment of the permissive H4K16ac histone mark in the Z-specific region in both tissues of females but not males, supporting the role of this histone mark in mediating dosage compensation of the Z chromosome.},
  author       = {Bett, Vincent K and Trejo Arellano, Minerva S and Vicoso, Beatriz},
  issn         = {1537-1719},
  journal      = {Molecular Biology and Evolution},
  number       = {5},
  publisher    = {Oxford University Press},
  title        = {{Chromatin landscape is associated with sex-biased expression and Drosophila-like dosage compensation of the Z chromosome in Artemia franciscana}},
  doi          = {10.1093/molbev/msaf085},
  volume       = {42},
  year         = {2025},
}

@misc{14705,
  abstract     = {Since the commercialization of brine shrimp (genus Artemia) in the 1950s, this lineage, and in particular the model species Artemia franciscana, has been the subject of extensive research. However, our understanding of the genetic mechanisms underlying various aspects of their reproductive biology, including sex determination, are still lacking. This is partly due to the scarcity of genomic resources for Artemia species and crustaceans in general. Here, we present a chromosome-level genome assembly of Artemia franciscana (Kellogg 1906), from the Great Salt Lake, USA. The genome is 1GB, and the majority of the genome (81%) is scaffolded into 21 linkage groups using a previously published high-density linkage map. We performed coverage and FST analyses using male and female genomic and transcriptomic reads to quantify the extent of differentiation between the Z and W chromosomes. Additionally, we quantified the expression levels in male and female heads and gonads and found further evidence for dosage compensation in this species.},
  author       = {Elkrewi, Marwan N},
  keywords     = {sex chromosome evolution, genome assembly, dosage compensation},
  publisher    = {Institute of Science and Technology Austria},
  title        = {{Data from "Chromosome-level assembly of Artemia franciscana sheds light on sex-chromosome differentiation"}},
  doi          = {10.15479/AT:ISTA:14705},
  year         = {2024},
}

