@article{12248,
  abstract     = {Eurasian brine shrimp (genus Artemia) have closely related sexual and asexual lineages of parthenogenetic females, which produce rare males at low frequencies. Although they are known to have ZW chromosomes, these are not well characterized, and it is unclear whether they are shared across the clade. Furthermore, the underlying genetic architecture of the transmission of asexuality, which can occur when rare males mate with closely related sexual females, is not well understood. We produced a chromosome-level assembly for the sexual Eurasian species Artemia sinica and characterized in detail the pair of sex chromosomes of this species. We combined this new assembly with short-read genomic data for the sexual species Artemia sp. Kazakhstan and several asexual lineages of Artemia parthenogenetica, allowing us to perform an in-depth characterization of sex-chromosome evolution across the genus. We identified a small differentiated region of the ZW pair that is shared by all sexual and asexual lineages, supporting the shared ancestry of the sex chromosomes. We also inferred that recombination suppression has spread to larger sections of the chromosome independently in the American and Eurasian lineages. Finally, we took advantage of a rare male, which we backcrossed to sexual females, to explore the genetic basis of asexuality. Our results suggest that parthenogenesis is likely partly controlled by a locus on the Z chromosome, highlighting the interplay between sex determination and asexuality.},
  author       = {Elkrewi, Marwan N and Khauratovich, Uladzislava and Toups, Melissa A and Bett, Vincent K and Mrnjavac, Andrea and Macon, Ariana and Fraisse, Christelle and Sax, Luca and Huylmans, Ann K and Hontoria, Francisco and Vicoso, Beatriz},
  issn         = {1943-2631},
  journal      = {Genetics},
  keywords     = {Genetics},
  number       = {2},
  publisher    = {Oxford University Press},
  title        = {{ZW sex-chromosome evolution and contagious parthenogenesis in Artemia brine shrimp}},
  doi          = {10.1093/genetics/iyac123},
  volume       = {222},
  year         = {2022},
}

@article{10166,
  abstract     = {While sexual reproduction is widespread among many taxa, asexual lineages have repeatedly evolved from sexual ancestors. Despite extensive research on the evolution of sex, it is still unclear whether this switch represents a major transition requiring major molecular reorganization, and how convergent the changes involved are. In this study, we investigated the phylogenetic relationship and patterns of gene expression of sexual and asexual lineages of Eurasian Artemia brine shrimp, to assess how gene expression patterns are affected by the transition to asexuality. We find only a few genes that are consistently associated with the evolution of asexuality, suggesting that this shift may not require an extensive overhauling of the meiotic machinery. While genes with sex-biased expression have high rates of expression divergence within Eurasian Artemia, neither female- nor male-biased genes appear to show unusual evolutionary patterns after sexuality is lost, contrary to theoretical expectations.},
  author       = {Huylmans, Ann K and Macon, Ariana and Hontoria, Francisco and Vicoso, Beatriz},
  issn         = {1471-2954},
  journal      = {Proceedings of the Royal Society B: Biological Sciences},
  keywords     = {asexual reproduction, parthenogenesis, sex-biased genes, sexual conflict, automixis, crustaceans},
  number       = {1959},
  publisher    = {The Royal Society},
  title        = {{Transitions to asexuality and evolution of gene expression in Artemia brine shrimp}},
  doi          = {10.1098/rspb.2021.1720},
  volume       = {288},
  year         = {2021},
}

@article{6418,
  abstract     = {Males and females of Artemia franciscana, a crustacean commonly used in the aquarium trade, are highly dimorphic. Sex is determined by a pair of ZW chromosomes, but the nature and extent of differentiation of these chromosomes is unknown. Here, we characterize the Z chromosome by detecting genomic regions that show lower genomic coverage in female than in male samples, and regions that harbor an excess of female-specific SNPs. We detect many Z-specific genes, which no longer have homologs on the W, but also Z-linked genes that appear to have diverged very recently from their existing W-linked homolog. We assess patterns of male and female expression in two tissues with extensive morphological dimorphism, gonads, and heads. In agreement with their morphology, sex-biased expression is common in both tissues. Interestingly, the Z chromosome is not enriched for sex-biased genes, and seems to in fact have a mechanism of dosage compensation that leads to equal expression in males and in females. Both of these patterns are contrary to most ZW systems studied so far, making A. franciscana an excellent model for investigating the interplay between the evolution of sexual dimorphism and dosage compensation, as well as Z chromosome evolution in general.},
  author       = {Huylmans, Ann K and Toups, Melissa A and Macon, Ariana and Gammerdinger, William J and Vicoso, Beatriz},
  issn         = {1759-6653},
  journal      = {Genome biology and evolution},
  number       = {4},
  pages        = {1033--1044},
  publisher    = {Oxford University Press},
  title        = {{Sex-biased gene expression and dosage compensation on the Artemia franciscana Z-chromosome}},
  doi          = {10.1093/gbe/evz053},
  volume       = {11},
  year         = {2019},
}

@article{190,
  abstract     = {The German cockroach, Blattella germanica, is a worldwide pest that infests buildings, including homes, restaurants, and hospitals, often living in unsanitary conditions. As a disease vector and producer of allergens, this species has major health and economic impacts on humans. Factors contributing to the success of the German cockroach include its resistance to a broad range of insecticides, immunity to many pathogens, and its ability, as an extreme generalist omnivore, to survive on most food sources. The recently published genome shows that B. germanica has an exceptionally high number of protein coding genes. In this study, we investigate the functions of the 93 significantly expanded gene families with the aim to better understand the success of B. germanica as a major pest despite such inhospitable conditions. We find major expansions in gene families with functions related to the detoxification of insecticides and allelochemicals, defense against pathogens, digestion, sensory perception, and gene regulation. These expansions might have allowed B. germanica to develop multiple resistance mechanisms to insecticides and pathogens, and enabled a broad, flexible diet, thus explaining its success in unsanitary conditions and under recurrent chemical control. The findings and resources presented here provide insights for better understanding molecular mechanisms that will facilitate more effective cockroach control.},
  author       = {Harrison, Mark and Arning, Nicolas and Kremer, Lucas and Ylla, Guillem and Belles, Xavier and Bornberg Bauer, Erich and Huylmans, Ann K and Jongepier, Evelien and Puilachs, Maria and Richards, Stephen and Schal, Coby},
  journal      = {Journal of Experimental Zoology Part B: Molecular and Developmental Evolution},
  pages        = {254--264},
  publisher    = {Wiley},
  title        = {{Expansions of key protein families in the German cockroach highlight the molecular basis of its remarkable success as a global indoor pest}},
  doi          = {10.1002/jez.b.22824},
  volume       = {330},
  year         = {2018},
}

@misc{9841,
  abstract     = {Around 150 million years ago, eusocial termites evolved from within the cockroaches, 50 million years before eusocial Hymenoptera, such as bees and ants, appeared. Here, we report the 2-Gb genome of the German cockroach, Blattella germanica, and the 1.3-Gb genome of the drywood termite Cryptotermes secundus. We show evolutionary signatures of termite eusociality by comparing the genomes and transcriptomes of three termites and the cockroach against the background of 16 other eusocial and non-eusocial insects. Dramatic adaptive changes in genes underlying the production and perception of pheromones confirm the importance of chemical communication in the termites. These are accompanied by major changes in gene regulation and the molecular evolution of caste determination. Many of these results parallel molecular mechanisms of eusocial evolution in Hymenoptera. However, the specific solutions are remarkably different, thus revealing a striking case of convergence in one of the major evolutionary transitions in biological complexity.},
  author       = {Harrison, Mark C. and Jongepier, Evelien and Robertson, Hugh M. and Arning, Nicolas and Bitard-Feildel, Tristan and Chao, Hsu and Childers, Christopher P. and Dinh, Huyen and Doddapaneni, Harshavardhan and Dugan, Shannon and Gowin, Johannes and Greiner, Carolin and Han, Yi and Hu, Haofu and Hughes, Daniel S. T. and Huylmans, Ann K and Kemena, Carsten and Kremer, Lukas P. M. and Lee, Sandra L. and Lopez-Ezquerra, Alberto and Mallet, Ludovic and Monroy-Kuhn, Jose M. and Moser, Annabell and Murali, Shwetha C. and Muzny, Donna M. and Otani, Saria and Piulachs, Maria-Dolors and Poelchau, Monica and Qu, Jiaxin and Schaub, Florentine and Wada-Katsumata, Ayako and Worley, Kim C. and Xie, Qiaolin and Ylla, Guillem and Poulsen, Michael and Gibbs, Richard A. and Schal, Coby and Richards, Stephen and Belles, Xavier and Korb, Judith and Bornberg-Bauer, Erich},
  publisher    = {Dryad},
  title        = {{Data from: Hemimetabolous genomes reveal molecular basis of termite eusociality}},
  doi          = {10.5061/dryad.51d4r},
  year         = {2018},
}

@article{448,
  abstract     = {Around 150 million years ago, eusocial termites evolved from within the cockroaches, 50 million years before eusocial Hymenoptera, such as bees and ants, appeared. Here, we report the 2-Gb genome of the German cockroach, Blattella germanica, and the 1.3-Gb genome of the drywood termite Cryptotermes secundus. We show evolutionary signatures of termite eusociality by comparing the genomes and transcriptomes of three termites and the cockroach against the background of 16 other eusocial and non-eusocial insects. Dramatic adaptive changes in genes underlying the production and perception of pheromones confirm the importance of chemical communication in the termites. These are accompanied by major changes in gene regulation and the molecular evolution of caste determination. Many of these results parallel molecular mechanisms of eusocial evolution in Hymenoptera. However, the specific solutions are remarkably different, thus revealing a striking case of convergence in one of the major evolutionary transitions in biological complexity.},
  author       = {Harrison, Mark and Jongepier, Evelien and Robertson, Hugh and Arning, Nicolas and Bitard Feildel, Tristan and Chao, Hsu and Childers, Christopher and Dinh, Huyen and Doddapaneni, Harshavardhan and Dugan, Shannon and Gowin, Johannes and Greiner, Carolin and Han, Yi and Hu, Haofu and Hughes, Daniel and Huylmans, Ann K and Kemena, Karsten and Kremer, Lukas and Lee, Sandra and López Ezquerra, Alberto and Mallet, Ludovic and Monroy Kuhn, Jose and Moser, Annabell and Murali, Shwetha and Muzny, Donna and Otani, Saria and Piulachs, Maria and Poelchau, Monica and Qu, Jiaxin and Schaub, Florentine and Wada Katsumata, Ayako and Worley, Kim and Xie, Qiaolin and Ylla, Guillem and Poulsen, Michael and Gibbs, Richard and Schal, Coby and Richards, Stephen and Belles, Xavier and Korb, Judith and Bornberg Bauer, Erich},
  journal      = {Nature Ecology and Evolution},
  number       = {3},
  pages        = {557--566},
  publisher    = {Springer Nature},
  title        = {{Hemimetabolous genomes reveal molecular basis of termite eusociality}},
  doi          = {10.1038/s41559-017-0459-1},
  volume       = {2},
  year         = {2018},
}

@article{945,
  abstract     = {While chromosome-wide dosage compensation of the X chromosome has been found in many species, studies in ZW clades have indicated that compensation of the Z is more localized and/or incomplete. In the ZW Lepidoptera, some species show complete compensation of the Z chromosome, while others lack full equalization, but what drives these inconsistencies is unclear. Here, we compare patterns of male and female gene expression on the Z chromosome of two closely related butterfly species, Papilio xuthus and Papilio machaon, and in multiple tissues of two moths species, Plodia interpunctella and Bombyx mori, which were previously found to differ in the extent to which they equalize Z-linked gene expression between the sexes. We find that, while some species and tissues seem to have incomplete dosage compensation, this is in fact due to the accumulation of male-biased genes and the depletion of female-biased genes on the Z chromosome. Once this is accounted for, the Z chromosome is fully compensated in all four species, through the up-regulation of Z expression in females and in some cases additional down-regulation in males. We further find that both sex-biased genes and Z-linked genes have increased rates of expression divergence in this clade, and that this can lead to fast shifts in patterns of gene expression even between closely related species. Taken together, these results show that the uneven distribution of sex-biased genes on sex chromosomes can confound conclusions about dosage compensation and that Z chromosome-wide dosage compensation is not only possible but ubiquitous among Lepidoptera.},
  author       = {Huylmans, Ann K and Macon, Ariana and Vicoso, Beatriz},
  issn         = {07374038},
  journal      = {Molecular Biology and Evolution},
  number       = {10},
  pages        = {2637 -- 2649},
  publisher    = {Oxford University Press},
  title        = {{Global dosage compensation is ubiquitous in Lepidoptera, but counteracted by the masculinization of the Z chromosome}},
  doi          = {10.1093/molbev/msx190},
  volume       = {34},
  year         = {2017},
}

@article{1019,
  abstract     = {As a consequence of its difference in copy number between males and females, the X chromosome is subject to unique evolutionary forces and gene regulatory mechanisms. Previous studies of Drosophila melanogaster have shown that the expression of X-linked, testis-specific reporter genes is suppressed in the male germline. However, it is not known whether this phenomenon is restricted to testis-expressed genes or if it is a more general property of genes with tissue-specific expression, which are also underrepresented on the X chromosome. To test this, we compared the expression of three tissue-specific reporter genes (ovary, accessory gland and Malpighian tubule) inserted at various autosomal and X-chromosomal locations. In contrast to testis-specific reporter genes, we found no reduction of X-linked expression in any of the other tissues. In accessory gland and Malpighian tubule, we detected higher expression of the X-linked reporter genes, which suggests that they are at least partially dosage compensated. We found no difference in the tissue-specificity of X-linked and autosomal reporter genes. These findings indicate that, in general, the X chromosome is not a detrimental environment for tissue-specific gene expression and that the suppression of X-linked expression is limited to the male germline.},
  author       = {Argyridou, Eliza and Huylmans, Ann K and Königer, Annabella and Parsch, John},
  issn         = {0018067X},
  journal      = {Heredity},
  number       = {1},
  pages        = {27 -- 34},
  publisher    = {Nature Publishing Group},
  title        = {{X-linkage is not a general inhibitor of tissue-specific gene expression in Drosophila melanogaster}},
  doi          = {10.1038/hdy.2017.12},
  volume       = {119},
  year         = {2017},
}

@misc{9861,
  abstract     = {As a consequence of its difference in copy number between males and females, the X chromosome is subject to unique evolutionary forces and gene regulatory mechanisms. Previous studies of Drosophila melanogaster have shown that the expression of X-linked, testis-specific reporter genes is suppressed in the male germline. However, it is not known whether this phenomenon is restricted to testis-expressed genes or if it is a more general property of genes with tissue-specific expression, which are also underrepresented on the X chromosome. To test this, we compared the expression of three tissue-specific reporter genes (ovary, accessory gland and Malpighian tubule) inserted at various autosomal and X-chromosomal locations. In contrast to testis-specific reporter genes, we found no reduction of X-linked expression in any of the other tissues. In accessory gland and Malpighian tubule, we detected higher expression of the X-linked reporter genes, which suggests that they are at least partially dosage compensated. We found no difference in the tissue-specificity of X-linked and autosomal reporter genes. These findings indicate that, in general, the X chromosome is not a detrimental environment for tissue-specific gene expression and that the suppression of X-linked expression is limited to the male germline.},
  author       = {Argyridou, Eliza and Huylmans, Ann K and Königer, Annabella and Parsch, John},
  publisher    = {Dryad},
  title        = {{Data from: X-linkage is not a general inhibitor of tissue-specific gene expression in Drosophila melanogaster}},
  doi          = {10.5061/dryad.02f6r},
  year         = {2017},
}

@article{1329,
  abstract     = {Daphnia species have become models for ecological genomics and exhibit interesting features, such as high phenotypic plasticity and a densely packed genome with many lineage-specific genes. They are also cyclic parthenogenetic, with alternating asexual and sexual cycles and environmental sex determination. Here, we present a de novo transcriptome assembly of over 32,000 D. galeata genes and use it to investigate gene expression in females and spontaneously produced males of two clonal lines derived from lakes in Germany and the Czech Republic. We find that only a low percentage (18%) of genes shows sex-biased expression and that there are many more female-biased gene (FBG) than male-biased gene (MBG). Furthermore, FBGs tend to be more conserved between species than MBGs in both sequence and expression. These patterns may be a consequence of cyclic parthenogenesis leading to a relaxation of purifying selection on MBGs. The two clonal lines show considerable differences in both number and identity of sex-biased genes, suggesting that they may have reproductive strategies differing in their investment in sexual reproduction. Orthologs of key genes in the sex determination and juvenile hormone pathways, which are thought to be important for the transition from asexual to sexual reproduction, are present in D. galeata and highly conserved among Daphnia species.},
  author       = {Huylmans, Ann K and López Ezquerra, Alberto and Parsch, John and Cordellier, Mathilde},
  journal      = {Genome Biology and Evolution},
  number       = {10},
  pages        = {3120 -- 3139},
  publisher    = {Oxford University Press},
  title        = {{De novo transcriptome assembly and sex-biased gene expression in the cyclical parthenogenetic Daphnia galeata}},
  doi          = {10.1093/gbe/evw221},
  volume       = {8},
  year         = {2016},
}

