{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T05:05:25Z","timestamp":1790053525727,"version":"4.0.1"},"reference-count":118,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,20]],"date-time":"2026-09-20T00:00:00Z","timestamp":1789862400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Ministry of Education and Science of the Republic of Kazakhstan","award":["AP23488097"],"award-info":[{"award-number":["AP23488097"]}],"id":[{"id":"https:\/\/ror.org\/03pj6ge82","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>Imprinting is an epigenetic mechanism found in many diploid organisms that enables discrimination between genetic material inherited from one or the other parent. It has been predominantly studied in mammals, where, in the adult, it affects the expression of a limited set of genes (in most higher mammals) or leads to paternal X chromosome inactivation in marsupials. DNA CpG methylation and heterochromatic epigenetic marks play a crucial role in these processes. Imprinting also exists in other organisms, where it can cause unusual parent-of-origin-dependent effects. Two classical examples are whole paternal genome inactivation in mealybug males and selective paternal X chromosome elimination in fungus gnats. Although these manifestations of imprinting are quite exotic, they share features in common with those observed in mammals, implying that, despite the differences, the core mechanism, the H3K9me3:HP1:H4K20me3 pathway, is evolutionarily conserved across animal phyla. Here, we discuss recent advances that provide further evidence supporting this concept.<\/jats:p>","DOI":"10.3390\/cells15181709","type":"journal-article","created":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T09:01:38Z","timestamp":1789981298000},"page":"1709","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Genomic Imprinting in Animals: Evolutionary Conservation with Adaptive Flexibility"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2315-4382","authenticated-orcid":false,"given":"Stepan N.","family":"Belyakin","sequence":"first","affiliation":[{"name":"Epigenetics Laboratory, Department of Natural Sciences, Novosibirsk State University, Pirogov Str. 2, 630090 Novosibirsk, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9571-0974","authenticated-orcid":false,"given":"Prim B.","family":"Singh","sequence":"additional","affiliation":[{"name":"Department of Biomedical Sciences, School of Medicine, Nazarbayev University, Kerey Zhanibek Khandar Street 5\/1, Astana 010000, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,20]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1429","DOI":"10.1093\/genetics\/45.10.1429","article-title":"The Controlling Element in Sex Chromosome Behavior in Sciara","volume":"45","author":"Crouse","year":"1960","journal-title":"Genetics"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1038\/s41586-024-07155-z","article-title":"Selfish conflict underlies RNA-mediated parent-of-origin effects","volume":"628","author":"Pliota","year":"2024","journal-title":"Nature"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1126\/science.145.3628.130","article-title":"Heterochromatic Chromosomes in the Coccids","volume":"145","author":"Brown","year":"1964","journal-title":"Science"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1038\/nrg3032","article-title":"Genomic imprinting: The emergence of an epigenetic paradigm","volume":"12","year":"2011","journal-title":"Nat. Rev. Genet."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1007\/s12038-016-9650-9","article-title":"Heterochromatin and the molecular mechanisms of \u2018parent-of-origin\u2019 effects in animals","volume":"41","author":"Singh","year":"2016","journal-title":"J. Biosci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1007\/s00412-018-00690-5","article-title":"Maternal regulation of chromosomal imprinting in animals","volume":"128","author":"Singh","year":"2019","journal-title":"Chromosoma"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Emmerson, R.A., and Mosher, R.A. (2026). Genomic Imprinting: Common Threads Uniting Diverse Biological Systems. Annu. Rev. Genet., 60.","DOI":"10.1146\/annurev-genet-012626-101213"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1016\/j.cell.2019.01.043","article-title":"Genomic Imprinting and Physiological Processes in Mammals","volume":"176","author":"Tucci","year":"2019","journal-title":"Cell"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1038\/315496a0","article-title":"Differential activity of maternally and paternally derived chromosome regions in mice","volume":"315","author":"Cattanach","year":"1985","journal-title":"Nature"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1016\/0092-8674(84)90313-1","article-title":"Completion of mouse embryogenesis requires both the maternal and paternal genomes","volume":"37","author":"McGrath","year":"1984","journal-title":"Cell"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"548","DOI":"10.1038\/308548a0","article-title":"Development of reconstituted mouse eggs suggests imprinting of the genome during gametogenesis","volume":"308","author":"Surani","year":"1984","journal-title":"Nature"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1098\/rsbl.2012.0666","article-title":"Facultative parthenogenesis discovered in wild vertebrates","volume":"8","author":"Booth","year":"2012","journal-title":"Biol. Lett."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1038\/nrg3766","article-title":"The role of genomic imprinting in biology and disease: An expanding view","volume":"15","author":"Peters","year":"2014","journal-title":"Nat. Rev. Genet."},{"key":"ref_14","first-page":"163","article-title":"DNA methylation and its biological significance","volume":"65","author":"Vaniushin","year":"1968","journal-title":"Usp. Sovrem. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"a018382","DOI":"10.1101\/cshperspect.a018382","article-title":"Genomic imprinting in mammals","volume":"6","author":"Barlow","year":"2014","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1038\/nrg3230","article-title":"Functions of DNA methylation: Islands, start sites, gene bodies and beyond","volume":"13","author":"Jones","year":"2012","journal-title":"Nat. Rev. Genet."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"a019133","DOI":"10.1101\/cshperspect.a019133","article-title":"DNA methylation in mammals","volume":"6","author":"Li","year":"2014","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"362","DOI":"10.1038\/366362a0","article-title":"Role for DNA methylation in genomic imprinting","volume":"366","author":"Li","year":"1993","journal-title":"Nature"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/0092-8674(85)90312-5","article-title":"A fraction of the mouse genome that is derived from islands of nonmethylated, CpG-rich DNA","volume":"40","author":"Bird","year":"1985","journal-title":"Cell"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1038\/ng1089","article-title":"Epigenetic regulation of gene expression: How the genome integrates intrinsic and environmental signals","volume":"33","author":"Jaenisch","year":"2003","journal-title":"Nat. Genet."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"9333","DOI":"10.1093\/nar\/12.24.9333","article-title":"Complete concordance between glucose-6-phosphate dehydrogenase activity and hypomethylation of 3\u2019 CpG clusters: Implications for X chromosome dosage compensation","volume":"12","author":"Wolf","year":"1984","journal-title":"Nucleic Acids Res."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1101\/sqb.2004.69.125","article-title":"Role of de novo DNA methyltransferases in initiation of genomic imprinting and X-chromosome inactivation","volume":"69","author":"Kaneda","year":"2004","journal-title":"Cold Spring Harb. Symp. Quant. Biol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1038\/nrg2295","article-title":"Epigenetic events in mammalian germ-cell development: Reprogramming and beyond","volume":"9","author":"Sasaki","year":"2008","journal-title":"Nat. Rev. Genet."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1101\/gr.130997.111","article-title":"Global profiling of DNA methylation erasure in mouse primordial germ cells","volume":"22","author":"Guibert","year":"2012","journal-title":"Genome Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"5285","DOI":"10.1074\/jbc.M108586200","article-title":"Maternal primary imprinting is established at a specific time for each gene throughout oocyte growth","volume":"277","author":"Obata","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1111\/j.1365-2443.2006.00943.x","article-title":"Oocyte growth-dependent progression of maternal imprinting in mice","volume":"11","author":"Hiura","year":"2006","journal-title":"Genes Cells"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"811","DOI":"10.1038\/ng.864","article-title":"Dynamic CpG island methylation landscape in oocytes and preimplantation embryos","volume":"43","author":"Smallwood","year":"2011","journal-title":"Nat. Genet."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Kobayashi, H., Sakurai, T., Imai, M., Takahashi, N., Fukuda, A., Yayoi, O., Sato, S., Nakabayashi, K., Hata, K., and Sotomaru, Y. (2012). Contribution of intragenic DNA methylation in mouse gametic DNA methylomes to establish oocyte-specific heritable marks. PLoS Genet., 8.","DOI":"10.1371\/journal.pgen.1002440"},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Shirane, K., Toh, H., Kobayashi, H., Miura, F., Chiba, H., Ito, T., Kono, T., and Sasaki, H. (2013). Mouse oocyte methylomes at base resolution reveal genome-wide accumulation of non-CpG methylation and role of DNA methyltransferases. PLoS Genet., 9.","DOI":"10.1371\/journal.pgen.1003439"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1008","DOI":"10.1016\/j.mod.2005.04.009","article-title":"Asymmetry in histone H3 variants and lysine methylation between paternal and maternal chromatin of the early mouse zygote","volume":"122","author":"Dieker","year":"2005","journal-title":"Mech. Dev."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2272","DOI":"10.1093\/hmg\/ddm179","article-title":"Role of the Dnmt3 family in de novo methylation of imprinted and repetitive sequences during male germ cell development in the mouse","volume":"16","author":"Kato","year":"2007","journal-title":"Hum. Mol. Genet."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2885","DOI":"10.1093\/hmg\/9.19.2885","article-title":"The H19 methylation imprint is erased and re-established differentially on the parental alleles during male germ cell development","volume":"9","author":"Davis","year":"2000","journal-title":"Hum. Mol. Genet."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1016\/j.molcel.2012.11.001","article-title":"The dynamics of genome-wide DNA methylation reprogramming in mouse primordial germ cells","volume":"48","author":"Seisenberger","year":"2012","journal-title":"Mol. Cell"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1101","DOI":"10.1038\/nature08829","article-title":"Genome-wide erasure of DNA methylation in mouse primordial germ cells is affected by AID deficiency","volume":"463","author":"Popp","year":"2010","journal-title":"Nature"},{"key":"ref_35","first-page":"455","article-title":"Dynamic distribution of the replacement histone variant H3.3 in the mouse oocyte and preimplantation embryos","volume":"50","author":"Bannister","year":"2006","journal-title":"Int. J. Dev. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1386","DOI":"10.1038\/nature04059","article-title":"The histone H3.3 chaperone HIRA is essential for chromatin assembly in the male pronucleus","volume":"437","author":"Loppin","year":"2005","journal-title":"Nature"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"811","DOI":"10.1242\/dev.061416","article-title":"Dynamic stage-specific changes in imprinted differentially methylated regions during early mammalian development and prevalence of non-CpG methylation in oocytes","volume":"138","author":"Tomizawa","year":"2011","journal-title":"Development"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1016\/j.molcel.2012.07.010","article-title":"Protection against de novo methylation is instrumental in maintaining parent-of-origin methylation inherited from the gametes","volume":"47","author":"Proudhon","year":"2012","journal-title":"Mol. Cell"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"20110336","DOI":"10.1098\/rstb.2011.0336","article-title":"New insights into establishment and maintenance of DNA methylation imprints in mammals","volume":"368","author":"Kelsey","year":"2013","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1038\/35000656","article-title":"Demethylation of the zygotic paternal genome","volume":"403","author":"Mayer","year":"2000","journal-title":"Nature"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1016\/S0960-9822(00)00448-6","article-title":"Active demethylation of the paternal genome in the mouse zygote","volume":"10","author":"Oswald","year":"2000","journal-title":"Curr. Biol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1530\/rep.1.00221","article-title":"Epigenetic reprogramming during early development in mammals","volume":"127","author":"Santos","year":"2004","journal-title":"Reproduction"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"829","DOI":"10.1016\/S0092-8674(01)00280-X","article-title":"Genomic imprinting disrupted by a maternal effect mutation in the Dnmt1 gene","volume":"104","author":"Howell","year":"2001","journal-title":"Cell"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"304","DOI":"10.1006\/dbio.2002.0628","article-title":"Dynamics of Dnmt1 methyltransferase expression and intracellular localization during oogenesis and preimplantation development","volume":"245","author":"Ratnam","year":"2002","journal-title":"Dev. Biol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1038\/nrm.2016.6","article-title":"Molecular features of cellular reprogramming and development","volume":"17","author":"Smith","year":"2016","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Singh, P.B., Belyakin, S.N., and Laktionov, P.P. (2020). Biology and Physics of Heterochromatin-Like Domains\/Complexes. Cells, 9.","DOI":"10.3390\/cells9081881"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1607","DOI":"10.1101\/gad.1667008","article-title":"Maternal and zygotic Dnmt1 are necessary and sufficient for the maintenance of DNA methylation imprints during preimplantation development","volume":"22","author":"Hirasawa","year":"2008","journal-title":"Genes Dev."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1016\/j.ydbio.2007.10.033","article-title":"Maintenance of genomic methylation patterns during preimplantation development requires the somatic form of DNA methyltransferase 1","volume":"313","author":"Kurihara","year":"2008","journal-title":"Dev. Biol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1101\/gad.234294.113","article-title":"DNA methylation dynamics during epigenetic reprogramming in the germline and preimplantation embryos","volume":"28","author":"Messerschmidt","year":"2014","journal-title":"Genes Dev."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1101\/sqb.2015.80.027441","article-title":"Erase-Maintain-Establish: Natural Reprogramming of the Mammalian Epigenome","volume":"80","author":"Leseva","year":"2016","journal-title":"Cold Spring Harb. Symp. Quant. Biol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1016\/j.devcel.2008.08.014","article-title":"A maternal-zygotic effect gene, Zfp57, maintains both maternal and paternal imprints","volume":"15","author":"Li","year":"2008","journal-title":"Dev. Cell"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1016\/j.molcel.2011.08.032","article-title":"In embryonic stem cells, ZFP57\/KAP1 recognize a methylated hexanucleotide to affect chromatin and DNA methylation of imprinting control regions","volume":"44","author":"Quenneville","year":"2011","journal-title":"Mol. Cell"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2374","DOI":"10.1101\/gad.202200.112","article-title":"An atomic model of Zfp57 recognition of CpG methylation within a specific DNA sequence","volume":"26","author":"Liu","year":"2012","journal-title":"Genes Dev."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1118","DOI":"10.1093\/nar\/gkv1059","article-title":"ZFP57 recognizes multiple and closely spaced sequence motif variants to maintain repressive epigenetic marks in mouse embryonic stem cells","volume":"44","author":"Anvar","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1186\/s13059-015-0672-7","article-title":"Allele-specific binding of ZFP57 in the epigenetic regulation of imprinted and non-imprinted monoallelic expression","volume":"16","author":"Strogantsev","year":"2015","journal-title":"Genome Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"6690","DOI":"10.1073\/pnas.1322273111","article-title":"Regulation of DNA methylation turnover at LTR retrotransposons and imprinted loci by the histone methyltransferase Setdb1","volume":"111","author":"Leung","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1159\/000547555","article-title":"Genomic Imprinting: Insights into Diverse Epigenetic Regulatory Mechanisms","volume":"166","author":"Pandey","year":"2026","journal-title":"Cytogenet. Genome Res."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1038\/39631","article-title":"Imprinted expression of the Igf2r gene depends on an intronic CpG island","volume":"389","author":"Wutz","year":"1997","journal-title":"Nature"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"810","DOI":"10.1038\/415810a","article-title":"The non-coding Air RNA is required for silencing autosomal imprinted genes","volume":"415","author":"Sleutels","year":"2002","journal-title":"Nature"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"20693","DOI":"10.1073\/pnas.1310201110","article-title":"H19 lncRNA controls gene expression of the Imprinted Gene Network by recruiting MBD1","volume":"110","author":"Monnier","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1038\/190372a0","article-title":"Gene action in the X-chromosome of the mouse (Mus musculus L.)","volume":"190","author":"Lyon","year":"1961","journal-title":"Nature"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"3478","DOI":"10.1038\/s44319-025-00499-1","article-title":"X chromosome inactivation in mammals: General principles and species-specific considerations","volume":"26","author":"Alfeghaly","year":"2025","journal-title":"EMBO Rep."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1038\/230231a0","article-title":"Late DNA replication in the paternally derived X chromosome of female kangaroos","volume":"230","author":"Sharman","year":"1971","journal-title":"Nature"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1038\/newbio230155a0","article-title":"Phosphoglycerate kinase polymorphism in kangaroos provides further evidence for paternal X inactivation","volume":"230","author":"Cooper","year":"1971","journal-title":"Nat. New. Biol."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Navarro-Cobos, M.J., and Brown, C.J. (2025). Human XIST: Origin and Divergence of a cis-Acting Silencing RNA. Noncoding RNA, 11.","DOI":"10.3390\/ncrna11030035"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"102446","DOI":"10.1016\/j.ceb.2024.102446","article-title":"Unleashing XIST from X-chromosome inactivation","volume":"92","author":"Morey","year":"2025","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1016\/S0959-437X(98)80090-7","article-title":"The role of Xist in X-inactivation","volume":"8","author":"Brockdorff","year":"1998","journal-title":"Curr. Opin. Genet. Dev."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1038\/nature11171","article-title":"Rsx is a metatherian RNA with Xist-like properties in X-chromosome inactivation","volume":"487","author":"Grant","year":"2012","journal-title":"Nature"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"17657","DOI":"10.1073\/pnas.0910322107","article-title":"Epigenetic modifications on X chromosomes in marsupial and monotreme mammals and implications for evolution of dosage compensation","volume":"107","author":"Rens","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Chaumeil, J., Waters, P.D., Koina, E., Gilbert, C., Robinson, T.J., and Graves, J.A. (2011). Evolution from XIST-independent to XIST-controlled X-chromosome inactivation: Epigenetic modifications in distantly related mammals. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0019040"},{"key":"ref_71","first-page":"20202244","article-title":"Koala methylomes reveal divergent and conserved DNA methylation signatures of X chromosome regulation","volume":"288","author":"Singh","year":"2021","journal-title":"Proc. Biol. Sci."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1101\/gr.161919.113","article-title":"Chromosome-wide profiling of X-chromosome inactivation and epigenetic states in fetal brain and placenta of the opossum, Monodelphis domestica","volume":"24","author":"Wang","year":"2014","journal-title":"Genome Res."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1093\/molbev\/msx297","article-title":"Landscape of DNA Methylation on the Marsupial X","volume":"35","author":"Waters","year":"2018","journal-title":"Mol. Biol. Evol."},{"key":"ref_74","first-page":"127","article-title":"Cytology of coccids (Coccoidea-Homoptera)","volume":"35","year":"1948","journal-title":"Adv. Genet."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1093\/genetics\/70.2.291","article-title":"RNA synthesis and heterochromatization in early development of a mealybug","volume":"70","author":"Sabour","year":"1972","journal-title":"Genetics"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"1072","DOI":"10.1242\/jcs.03412","article-title":"Epigenetic regulation of facultative heterochromatinisation in Planococcus citri via the Me(3)K9H3-HP1-Me(3)K20H4 pathway","volume":"120","author":"Bongiorni","year":"2007","journal-title":"J. Cell Sci."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1007\/BF00284926","article-title":"Cytogenetics of the mealybug Planococcus citri (Risso) (Homoptera: Coccoidea): Genetic markers, lethals, and chromosome rearrangements","volume":"28","author":"Brown","year":"1969","journal-title":"Chromosoma"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1093\/genetics\/46.8.983","article-title":"Radiation Analysis of a Lecanoid Genetic System","volume":"46","author":"Brown","year":"1961","journal-title":"Genetics"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1007\/s00412-002-0182-8","article-title":"Heterochromatin, HP1 and methylation at lysine 9 of histone H3 in animals","volume":"111","author":"Cowell","year":"2002","journal-title":"Chromosoma"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"2491","DOI":"10.1242\/jcs.01238","article-title":"Heterochromatin and tri-methylated lysine 20 of histone H4 in animals","volume":"117","author":"Kourmouli","year":"2004","journal-title":"J. Cell Sci."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1007\/BF00336754","article-title":"Meiotic parthenogenesis and heterochromatization in a soft scale, pulvinaria hydrangeae (Coccoidea: Homoptera)","volume":"14","author":"Nur","year":"1963","journal-title":"Chromosoma"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1093\/icb\/11.2.301","article-title":"Parthenogenesis in Coccids (Homoptera)","volume":"11","author":"Nur","year":"1971","journal-title":"Am. Zool."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1007\/BF00326174","article-title":"Diploid arrhenotoky and automictic thelytoky in soft scale insects (Lecaniidae: Coccoidea: Homoptera)","volume":"39","author":"Nur","year":"1972","journal-title":"Chromosoma"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1038\/253165a0","article-title":"Chromosome imprinting and the mammalian X chromosome","volume":"253","author":"Chandra","year":"1975","journal-title":"Nature"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1471","DOI":"10.1093\/genetics\/151.4.1471","article-title":"The relationship between DNA methylation and chromosome imprinting in the coccid Planococcus citri","volume":"151","author":"Bongiorni","year":"1999","journal-title":"Genetics"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"5687","DOI":"10.1111\/mec.15842","article-title":"Sex-specific expression and DNA methylation in a species with extreme sexual dimorphism and paternal genome elimination","volume":"30","author":"Bain","year":"2021","journal-title":"Mol. Ecol."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"19466","DOI":"10.1074\/jbc.RA118.004612","article-title":"BAH domains and a histone-like motif in DNA methyltransferase 1 (DNMT1) regulate de novo and maintenance methylation in vivo","volume":"293","author":"Yarychkivska","year":"2018","journal-title":"J. Biol. Chem."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"594","DOI":"10.1038\/s41594-021-00603-8","article-title":"Dnmt1 has de novo activity targeted to transposable elements","volume":"28","author":"Haggerty","year":"2021","journal-title":"Nat. Struct. Mol. Biol."},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Omar, M.A.A., Li, M., Liu, F., He, K., Qasim, M., Xiao, H., Jiang, M., and Li, F. (2020). The Roles of DNA Methyltransferases 1 (DNMT1) in Regulating Sexual Dimorphism in the Cotton Mealybug, Phenacoccus solenopsis. Insects, 11.","DOI":"10.3390\/insects11020121"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1242\/jcs.101.2.463","article-title":"Cloning and expression of Drosophila HP1 homologs from a mealybug, Planococcus citri","volume":"101","author":"Epstein","year":"1992","journal-title":"J. Cell Sci."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1007\/s00412-023-00809-3","article-title":"H3K9 and H4K20 methyltransferases are directly involved in the heterochromatinization of the paternal chromosomes in male Planococcus citri embryos","volume":"132","author":"Osipov","year":"2023","journal-title":"Chromosoma"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1007\/978-3-540-39838-7_2","article-title":"Unusual chromosome movements in sciarid flies","volume":"13","author":"Gerbi","year":"1986","journal-title":"Results Probl. Cell Differ."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1086\/280803","article-title":"Chromosome behavior, inheritance and sex determination in Sciara","volume":"72","author":"Metz","year":"1938","journal-title":"Am. Nat."},{"key":"ref_94","doi-asserted-by":"crossref","unstructured":"Singh, P.B., and Belyakin, S.N. (2018). L Chromosome Behaviour and Chromosomal Imprinting in Sciara Coprophila. Genes, 9.","DOI":"10.3390\/genes9090440"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1002\/1521-1878(200103)23:3<242::AID-BIES1034>3.0.CO;2-P","article-title":"Chromosome elimination in sciarid flies","volume":"23","author":"Goday","year":"2001","journal-title":"Bioessays"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"3775","DOI":"10.1242\/dev.122.12.3775","article-title":"Incomplete sister chromatid separation is the mechanism of programmed chromosome elimination during early Sciara coprophila embryogenesis","volume":"122","author":"Sullivan","year":"1996","journal-title":"Development"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1002\/jmor.1050680307","article-title":"Chromosome behavior in the germ cells and development of the gonads of Sciara ocellaris","volume":"68","author":"Berry","year":"1941","journal-title":"J. Morph."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"msad148","DOI":"10.1093\/molbev\/msad148","article-title":"Recent Evolution of a Maternally Acting Sex-Determining Supergene in a Fly with Single-Sex Broods","volume":"40","author":"Baird","year":"2023","journal-title":"Mol. Biol. Evol."},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Posukh, O.V., Shloma, V.V., Skrypnik, P.A., Maksimov, D.A., Antoshina, P.A., Kalashnikova, D.A., Nurislamov, A., Lukyanchikova, V.A., Torgunakov, N., and Battulin, N.R. (2025). DNA and Histone Modifications Identify a Putative Controlling Element (CE) on the X Chromosome of Sciara coprophila. Cells, 14.","DOI":"10.3390\/cells14161243"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1007\/BF00294938","article-title":"Localization of ribosomal DNA within the proximal X heterochromatin of Sciara coprophila (Diptera, Sciaridae)","volume":"64","author":"Crouse","year":"1977","journal-title":"Chromosoma"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1007\/BF00328650","article-title":"The nature of the influence of x-translocations on sex of progeny in Sciara coprophila","volume":"11","author":"Crouse","year":"1960","journal-title":"Chromosoma"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1007\/BF00292274","article-title":"X heterochromatin subdivision and cytogenetic analysis in Sciara coprophila (diptera, sciaridae)","volume":"74","author":"Crouse","year":"1979","journal-title":"Chromosoma"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"1715","DOI":"10.1242\/jcs.083022","article-title":"Histone H3 phosphorylation and non-disjunction of the maternal X chromosome during male meiosis in sciarid flies","volume":"124","author":"Escriba","year":"2011","journal-title":"J. Cell Sci."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1126\/science.1566060","article-title":"Maternal-effect selfish genes in flour beetles","volume":"256","author":"Beeman","year":"1992","journal-title":"Science"},{"key":"ref_105","doi-asserted-by":"crossref","unstructured":"Seidel, H.S., Ailion, M., Li, J., van Oudenaarden, A., Rockman, M.V., and Kruglyak, L. (2011). A novel sperm-delivered toxin causes late-stage embryo lethality and transmission ratio distortion in C. elegans. PLoS Biol., 9.","DOI":"10.1371\/journal.pbio.1001115"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"1051","DOI":"10.1126\/science.aan0621","article-title":"A maternal-effect selfish genetic element in Caenorhabditis elegans","volume":"356","author":"Burga","year":"2017","journal-title":"Science"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"990","DOI":"10.1016\/j.cub.2020.12.013","article-title":"Ubiquitous Selfish Toxin-Antidote Elements in Caenorhabditis Species","volume":"31","author":"Pliota","year":"2021","journal-title":"Curr. Biol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"589","DOI":"10.1126\/science.1151107","article-title":"Widespread genetic incompatibility in C. elegans maintained by balancing selection","volume":"319","author":"Seidel","year":"2008","journal-title":"Science"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"RP93194","DOI":"10.7554\/eLife.93194","article-title":"Evolutionary adaptation of an HP1-protein chromodomain integrates chromatin and DNA sequence signals","volume":"13","author":"Baumgartner","year":"2024","journal-title":"eLife"},{"key":"ref_110","first-page":"585024","article-title":"Epigenetic mechanisms of genomic imprinting: Common themes in the regulation of imprinted regions in mammals, plants, and insects","volume":"2012","author":"Macdonald","year":"2012","journal-title":"Genet. Res. Int."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"102764","DOI":"10.1016\/j.pbi.2025.102764","article-title":"Rapid origin and turnover of genomic imprinting by transposable elements","volume":"87","author":"Kohler","year":"2025","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1080\/10409238.2017.1279119","article-title":"DNA methylation and imprinting in plants: Machinery and mechanisms","volume":"52","author":"Satyaki","year":"2017","journal-title":"Crit. Rev. Biochem. Mol. Biol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1038\/nature05914","article-title":"Transcription and RNA interference in the formation of heterochromatin","volume":"447","author":"Grewal","year":"2007","journal-title":"Nature"},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"MacDonald, W.A., and Mann, M.R.W. (2020). Long noncoding RNA functionality in imprinted domain regulation. PLoS Genet., 16.","DOI":"10.1371\/journal.pgen.1008930"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1038\/s41576-020-0245-9","article-title":"Maternal H3K27me3-dependent autosomal and X chromosome imprinting","volume":"21","author":"Chen","year":"2020","journal-title":"Nat. Rev. Genet."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1038\/nature23262","article-title":"Maternal H3K27me3 controls DNA methylation-independent imprinting","volume":"547","author":"Inoue","year":"2017","journal-title":"Nature"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"eaay7246","DOI":"10.1126\/sciadv.aay7246","article-title":"Allelic H3K27me3 to allelic DNA methylation switch maintains noncanonical imprinting in extraembryonic cells","volume":"5","author":"Chen","year":"2019","journal-title":"Sci. Adv."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1186\/s13059-019-1833-x","article-title":"Endogenous retroviral insertions drive non-canonical imprinting in extra-embryonic tissues","volume":"20","author":"Hanna","year":"2019","journal-title":"Genome Biol."}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/18\/1709\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T04:41:37Z","timestamp":1790052097000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/18\/1709"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,20]]},"references-count":118,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["cells15181709"],"URL":"https:\/\/doi.org\/10.3390\/cells15181709","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,20]]}}}