329:: the mature oocyte has an intermediate level of DNA methylation (72%), instead the sperm has high level of DNA methylation (86%). Demethylation in paternal genome occurs quickly after fertilisation, whereas the maternal genome is quite resistant at the demethylation process at this stage. Maternal different methylated regions (DMRs) are more resistant to the preimplantation demethylation wave.
400:
exceeding 30% differences in 5% of the loci. The stochastic switching occurred at thousands of heterozygous regulatory loci that were bound to transcription factors. The intermediate methylation state is referred to the relative frequencies between methylated and unmethylated epialleles. The epiallele frequency variations are correlated with the allele affinity for transcription factors.
22:
540:(ncRNA) gene silencing involves various types of non-coding RNAs, such as microRNAs (miRNAs), long non-coding RNAs (lncRNAs), and small interfering RNAs (siRNAs). These RNA molecules can modulate gene expression by various mechanisms, including mRNA degradation, inhibition of translation, and chromatin remodeling.
650:
Further proofs towards a role in genetic modulation and transcription regulation refers to the great conservation of the boundary pattern across mammalian evolution, with a dynamic range of small diversities inside different cell types, suggesting that these topological domains take part in cell-type
407:
Haplotype reconstruction strategy is used to trace chromatin chemical modifications (using ChIP-seq) in a variety of human tissues. Haplotype-resolved epigenomic maps can trace allelic biases in chromatin configuration. A substantial variation among different tissues and individuals is observed. This
586:
of the cell. They are formed by regions of chromatin, sized from 100 kilobases up to megabases, which highly self-interact. The domains are linked by other genomic regions, which, based on their size, are either called “topological boundary regions” or “unorganized chromatin”. These boundary regions
246:
CoRSIVs can have a useful application: measurements of CoRSIV methylation in one tissue can provide some information about epigenetic regulation in other tissues, indeed we can predict the expression of associated genes because systemic epigenetic variants are generally consistent in all tissues and
659:
The 4D Nucleome project aims to realize a 3D maps of mammalian genomes in order to develop predictive models to correlate epigenomic modifications with genetic variation. In particular the goal is to link genetic and epigenomic modifications with the enhancers and promoters which they interact with
264:
polymorphisms in humans suggests that such effects are highly deleterious. Indeed, trans-acting factors are expected to be caused by mutations in chromatin control genes or other highly pleiotropic regulators. If trans-acting variants do exist in human populations, they probably segregate as rare
399:
was aimed to construct the maps of allelic imbalances in DNA methylation, gene transcription, and also of histone modifications. 36 cell and tissue types from 13 participant donors were used to examine 71 epigenomes. The results of WGBS tested on 49 methylomes revealed CpG methylation imbalances
742:
has been widely supported since it was initially developed the 1980s. In recent decades, evidence has accumulated supporting the additional idea that DNA damage and repair elicit widespread epigenome alterations that also contribute to aging (e.g.). Such epigenome changes include age-related
207:
ariation in DNA methylation. They span only 0.1% of the human genome, so they are very rare; they can be inter-correlated over long genomic distances (>50 kbp). CoRSIVs are also associated with genes involved in a lot of human disorders, including tumors, mental disorders and cardiovascular
255:
Quantification of the heritable basis underlying population epigenomic variation is also important to delineate its cis- and trans-regulatory architecture. In particular, most studies state that inter-individual differences in DNA methylation are mainly determined by cis-regulatory sequence
514:
Histone acetylation neutralizes the positive charge on histones. This weakens the electrostatic attraction to negatively charged DNA and causes unwinding of DNA from histones, making the DNA more accessible to the transcriptional machinery and hence resulting in transcriptional activation.
384:(WGBS) is used to explore sequence-dependent allele-specific methylation (SD-ASM) at a single-chromosome resolution level and comprehensive whole-genome coverage. The results of WGBS tested on 49 methylomes revealed CpG methylation imbalances exceeding 30% differences in 5% of the loci.
211:
Most of the CoRSIVs are only 200 – 300 bp long and include 5–10 CpG dinucleotides, the largest span several kb and involve hundreds of CpGs. These regions tend to occur in clusters and the two genomic areas of high CoRSIV density are observed at the major histocompatibility
281:
Anyway, differential expression concerns only a slight number of methylated genes: only one fifth of genes with CpG methylation shows variable expression according to their methylation state. It is important to notice that methylation is not the only factor affecting
362:
to the 8-cell stage and then increase. Parental allele-specific analysis shows that paternal genome becomes more open than the maternal genome from the late zygote stage to the 4-cell stage, which may reflect decondensation of the paternal genome with replacement of
598:
process. Inside these domains, the chromatin shows to be well tangled, while in the boundary regions chromatin interactions are far less present. These boundary areas in particular show some peculiarity that determine the functions of all the topological domains.
403:
The analysis of the study suggests that human epigenome in average covers approximately 200 adverse SD-ASM variants. The sensitivity of the genes with tissue-specific expression patterns gives the opportunity for the evolutionary innovation in gene regulation.
379:
DNA methylation imbalances between homologous chromosomes show sequence-dependent behavior. Difference in the methylation state of neighboring cytosines on the same chromosome occurs due to the difference in DNA sequence between the chromosomes. Whole-genome
548:
During the last few years, several methods have been developed to study the structural and consequently the functional modifications of chromatin. The first project that used epigenomic profiling to identify regulatory elements in the human genome was
387:
On the sites of gene regulatory loci bound by transcription factors the random switching between methylated and unmethylated states of DNA was observed. This is also referred as stochastic switching and it is linked to selective buffering of
793:
is to generate human reference epigenomes from normal, healthy individuals across a large variety of cell lines, primary cells, and primary tissues. Data produced by the project, which can be browsed and downloaded from the
273:
DNA methylation (in particular in CpG regions) is able to affect gene expression: hypermethylated regions tend to be differentially expressed. In fact, people with a similar methylation profile tend to also have the same
763:. Advances in sequencing technology now allow for assaying genome-wide epigenomic states by multiple molecular methodologies. Micro- and nanoscale devices have been constructed or proposed to investigate the epigenome.
208:
diseases. It has been observed that disease-associated CpG sites are 37% enriched in CoRSIVs compared to control regions and 53% enriched in CoRSIVs relative to tDMRs (tissue specific
Differentially Methylated Regions).
2352:
632:
genes are particularly abundant in boundary regions, denoting that those areas have a prolific transcriptional activity, thanks to their structural characteristics, different from other topological regions.
587:
separate the topological domains from heterochromatin, and prevent the amplification of the latter. Topological domains are diffused in mammalian, although similar genome partitions were identified also in
174:
methylation among individuals is about 42%. On the contrary, epigenetic profile (including methylation profile) of each individual is constant over the course of a year, reflecting the constancy of our
617:
Secondly, boundary regions block heterochromatin spreading, thus preventing the loss of useful genetic informations. This information derives from the observation that the heterochromatin mark
877:
Reference epigenomes for healthy individuals will enable the second goal of the
Roadmap Epigenomics Project, which is to examine epigenomic differences that occur in disease states such as
827:), and Methylation-sensitive Restriction Enzyme Sequencing (MRE-Seq) identify DNA methylation across portions of the genome at varying levels of resolution down to basepair level.
231:
regions) and contain many transposable elements, but few CpG islands (CGI) and transcription factor binding sites. CoRSIVs are under-represented in the proximity of genes, in
2349:
358:. Stage-specific proximal and distal regions with accessible chromatin regions were identified. Global chromatin accessibility is found to gradually decrease from the
278:. Moreover, one key observation from human methylation is that most functionally relevant changes in CpG methylation occur in regulatory elements, such as enhancers.
93:, enabling cells with the same genetic code to perform different functions. The human epigenome is dynamic and can be influenced by environmental factors such as
2384:"Multilayer-omics analyses of human cancers: exploration of biomarkers and drug targets based on the activities of the International Human Epigenome Consortium"
438:. These modifications can either activate or repress gene expression by altering chromatin structure and accessibility of the DNA to transcriptional machinery.
667:
Hi-C is an experimental method used to map the connections between DNA fragments in three-dimensional space on a genome-wide scale. This technique combines
260:, probably involving mutations in TFBSs (Transcription Factor Binding Sites) with downstream consequences on local chromatin environment. The sparsity of
112:. Unlike the underlying genome, which remains largely static within an individual, the epigenome can be dynamically altered by environmental conditions.
770:(IHEC). IHEC members aim to generate at least 1,000 reference (baseline) human epigenomes from different types of normal and disease-related human
1589:"Conserved, developmentally regulated mechanism couples chromosomal looping and heterochromatin barrier activity at the homeobox gene A locus"
647:. In the recent years, those sequences were referred to alter binding site of CTCF, thus interfering with expression of some genomic areas.
553:(Encyclopedia of DNA Elements) that focused on profiling histone modifications on cell lines. A few years later ENCODE was included in the
179:
and metabolic traits. Methylation profile, in particular, is quite stable in a 12-month period and appears to change more over decades.
314:. This is a consequence of active DNA demethylation at this stage. But global demethylation is not an irreversible process, in fact
767:
554:
54:
798:, fall into five types that assay different aspects of the epigenome and outcomes of epigenomic states (such as gene expression):
710:
modification patterns. The aberrant epigenetic landscape of the cancer cell is characterized by a global genomic hypomethylation,
159:(5mC). This epigenetic mark is widely conserved and plays major roles in the regulation of gene expression, in the silencing of
2484:"Comparison of sequencing-based methods to profile DNA methylation and identification of monoallelic epigenetic modifications"
441:
The epigenetic profiles of human tissues reveals the following distinct histone modifications in different functional areas:
824:
641:
2539:
2333:
900:
731:
579:
529:
Can lead to activation or repression of gene expression depending on the specific amino acids that are methylated.
213:
265:
alleles or originate from somatic mutations and present with clinical phenotypes, as is the case in many cancers.
108:
The epigenome is involved in regulating gene expression, development, tissue differentiation, and suppression of
2581:
2062:
1795:"Waves of retrotransposon expansion remodel genome organization and CTCF binding in multiple mammalian lineages"
786:
392:
against mutations and genetic diseases. Only rare genetic variants show the stochastic type of gene regulation.
2572:
283:
594:
Topological domains in humans, like in other mammalians, have many functions regarding gene expression and
155:
DNA functionally interacts with a variety of epigenetic marks, such as cytosine methylation, also known as
836:
636:
Finally, in the border areas of the topological domains and their surroundings there is an enrichment of
1089:"Omic personality: implications of stable transcript and methylation profiles for personalized medicine"
1041:
Taudt A, Colomé-Tatché M, Johannes F (June 2016). "Genetic sources of population epigenomic variation".
595:
811:) identifies genome wide patterns of histone modifications using antibodies against the modifications.
318:
methylation occurring from the early to mid-pronuclear stage and from the 4-cell to the 8-cell stage.
606:
regions and barrier elements, both of which function as inhibitors of further transcription from the
408:
allows the deeper understanding of cis-regulatory relationships between genes and control sequences.
2013:
Yang JH, Hayano M, Griffin PT, Amorim JA, Bonkowski MS, Apostolides JK, et al. (January 2023).
1194:
Waterland RA, Michels KB (2007). "Epigenetic epidemiology of the developmental origins hypothesis".
560:
The structural modifications that these projects aim to study can be divided into five main groups:
1844:"High-resolution genetic mapping of putative causal interactions between regions of open chromatin"
1793:
Schmidt D, Schwalie PC, Wilson MD, Ballester B, Gonçalves A, Kutter C, et al. (January 2012).
1486:
Sexton T, Yaffe E, Kenigsberg E, Bantignies F, Leblanc B, Hoichman M, et al. (February 2012).
90:
1282:"Allele-specific epigenome maps reveal sequence-dependent stochastic switching at regulatory loci"
735:
389:
164:
1280:
Onuchic V, Lurie E, Carrero I, Pawliczek P, Patel RY, Rozowsky J, et al. (September 2018).
878:
257:
2435:"Genome-wide chromatin state transitions associated with developmental and environmental cues"
1744:
Ebersole T, Kim JH, Samoshkin A, Kouprina N, Pavlicek A, White RJ, et al. (August 2011).
823:, Reduced Representation Bisulfite-Seq (RRBS), Methylated DNA Immunoprecipitation Sequencing (
610:
enzyme. Such elements are characterized by the massive presence of insulator binding proteins
1143:
Gunasekara CJ, Scott CA, Laritsky E, Baker MS, MacKay H, Duryea JD, et al. (June 2019).
668:
109:
86:
2601:
2205:
2149:
1600:
1543:
1532:"Topological domains in mammalian genomes identified by analysis of chromatin interactions"
1488:"Three-dimensional folding and functional organization principles of the Drosophila genome"
1443:
Stricker SH, Köferle A, Beck S (January 2017). "From profiles to function in epigenomics".
1345:
820:
766:
An international effort to assay reference epigenomes commenced in 2010 in the form of the
603:
417:
381:
160:
149:
74:
2482:
Harris RA, Wang T, Coarfa C, Nagarajan RP, Hong C, Downey SL, et al. (October 2010).
2138:"Micro- and nanoscale devices for the investigation of epigenetics and chromatin dynamics"
1332:
Leung D, Jung I, Rajagopal N, Schmitt A, Selvaraj S, Lee AY, et al. (February 2015).
1207:
722:
code for critical genes and a global loss of monoacetylated and trimethylated histone H4.
8:
866:
524:
509:
240:
236:
2586:
2209:
2153:
2039:
2014:
1697:"Regulating RNA polymerase pausing and transcription elongation in embryonic stem cells"
1604:
1547:
1349:
2508:
2483:
2459:
2434:
2410:
2383:
2308:
2283:
2170:
2137:
2113:
2088:
1917:
1892:
1868:
1843:
1819:
1794:
1770:
1745:
1721:
1696:
1672:
1647:
1623:
1588:
1564:
1531:
1468:
1366:
1333:
1306:
1281:
1171:
1144:
1115:
1088:
1066:
672:
1940:
Gensler HL, Bernstein H (September 1981). "DNA damage as the primary cause of aging".
678:
This kind of studies are currently limited by the lack or unavailability of raw data.
2513:
2464:
2415:
2313:
2264:
2223:
2175:
2118:
2044:
1995:
1978:
Siametis A, Niotis G, Garinis GA (April 2021). "DNA Damage and the Aging
Epigenome".
1957:
1922:
1873:
1824:
1775:
1726:
1677:
1628:
1569:
1509:
1460:
1420:
1415:
1390:
1371:
1311:
1259:
1211:
1176:
1120:
1058:
1020:
988:
953:
895:
795:
625:
224:
25:
The function of DNA strands (yellow) alters depending on how it is organized around
2503:
2495:
2454:
2446:
2433:
Zhu J, Adli M, Zou JY, Verstappen G, Coyne M, Zhang X, et al. (January 2013).
2405:
2395:
2303:
2295:
2254:
2213:
2165:
2157:
2108:
2100:
2034:
2026:
1987:
1949:
1912:
1904:
1863:
1855:
1814:
1806:
1765:
1757:
1716:
1708:
1667:
1659:
1618:
1608:
1559:
1551:
1499:
1472:
1452:
1410:
1406:
1402:
1361:
1353:
1301:
1293:
1249:
1203:
1166:
1156:
1110:
1100:
1050:
980:
943:
910:
333:
156:
98:
94:
2544:
1646:
Hawkins RD, Hon GC, Lee LK, Ngo Q, Lister R, Pelizzola M, et al. (May 2010).
1070:
759:
aims to identify and catalogue
Methylation Variable Positions (MVPs) in the human
664:
and pathways as new candidates for functional analysis and therapeutic targeting.
2576:
2356:
2337:
1648:"Distinct epigenomic landscapes of pluripotent and lineage-committed human cells"
1254:
1238:"Human Germline Cell Development: from the Perspective of Single-Cell Sequencing"
1237:
905:
847:
815:
808:
790:
739:
703:
644:
427:
422:
Post-translational modifications of histone proteins, which include methylation,
307:
232:
135:
70:
50:
1012:
2450:
2030:
1908:
1810:
1663:
1593:
Proceedings of the
National Academy of Sciences of the United States of America
1504:
1487:
948:
931:
856:
607:
537:
431:
295:
145:
2104:
1991:
1859:
1161:
1105:
170:
Individuals differ with their epigenetic profile, for example the variance in
2595:
2400:
1746:"tRNA genes protect a reporter gene from epigenetic silencing in mouse cells"
303:
299:
275:
1695:
Min IM, Waterfall JJ, Core LJ, Munroe RJ, Schimenti J, Lis JT (April 2011).
1613:
1530:
Dixon JR, Selvaraj S, Yue F, Kim A, Li Y, Shen Y, et al. (April 2012).
1334:"Integrative analysis of haplotype-resolved epigenomes across human tissues"
1297:
1145:"A genomic atlas of systemic interindividual epigenetic variation in humans"
1087:
Tabassum R, Sivadas A, Agrawal V, Tian H, Arafat D, Gibson G (August 2015).
2517:
2468:
2419:
2330:
2317:
2299:
2268:
2227:
2179:
2161:
2122:
2048:
1999:
1926:
1877:
1828:
1779:
1761:
1730:
1681:
1632:
1573:
1513:
1464:
1424:
1375:
1315:
1263:
1215:
1180:
1124:
1062:
1024:
992:
957:
843:) uses the DNase I enzyme to find open or accessible regions in the genome.
629:
341:
261:
217:
1961:
344:(MII) stage. Non-CpG methylation continues to accumulate in these stages.
1456:
890:
691:
661:
637:
435:
423:
337:
243:. They are also usually not present in highly conserved genomic regions.
228:
121:
1712:
1555:
1357:
1054:
350:
accessibility in germline was evaluated by different approaches, like sc
298:
experiments that in human preimplantation embryos there is a global DNA
771:
711:
588:
564:
311:
984:
268:
2499:
2066:
840:
831:
783:
743:
changes in the patterns of DNA methylation and histone modification.
364:
355:
347:
176:
58:
57:. Changes to the epigenome can result in changes to the structure of
2564:
2369:
2259:
2242:
2218:
2193:
220:
and at the pericentromeric region on the long arm of chromosome 20.
1953:
971:
Delcuve GP, Rastegar M, Davie JR (May 2009). "Epigenetic control".
870:
654:
476:
471:
351:
171:
141:
2569:
557:(IHEC), which aims to coordinate international epigenome studies.
53:; these changes can be passed down to an organism's offspring via
852:
803:
719:
707:
618:
495:
488:
483:
466:
461:
374:
368:
82:
78:
46:
34:
26:
2534:
1485:
760:
695:
583:
550:
359:
322:
62:
2015:"Loss of epigenetic information as a cause of mammalian aging"
1792:
715:
699:
326:
140:
Addition of a methyl group to the DNA molecule, typically at
102:
2549:
1743:
1279:
1142:
41:
of an organism is the collection of chemical changes to its
21:
1391:"Acetylation of histones and transcription-related factors"
611:
2554:
2284:"Perspectives of international human epigenome consortium"
1086:
1040:
2559:
2481:
1331:
42:
1893:"Principles of Chromosome Architecture Revealed by Hi-C"
869:
identifies expression of small noncoding RNA, primarily
574:
250:
2012:
929:
2560:
2350:"France: Human epigenome consortium takes first steps"
1841:
1694:
660:
in three-dimensional space, thus discovering gene-set
621:
sequences clearly interrupts near boundary sequences.
49:
proteins that affects when, where, and how the DNA is
1977:
930:
Bernstein BE, Meissner A, Lander ES (February 2007).
2432:
1010:
970:
2080:
1842:Kumasaka N, Knights AJ, Gaffney DJ (January 2019).
1529:
1442:
859:
identify expression levels or protein coding genes.
269:
Correlation between methylation and gene expression
567:occupancy to detect regions with regulatory genes;
321:The percentage of DNA methylation is different in
152:and other proteins necessary for gene expression.
2593:
2565:Human Epigenome Browser at Washington University
2135:
1939:
1645:
1193:
655:Correlation between methylation and 3D structure
532:
2535:Reference Epigenome Mapping Consortium Homepage
1973:
1971:
1019:. Treasure Island (FL): StatPearls Publishing.
1011:Al Aboud NM, Tupper C, Jialal I (August 2023).
1006:
1004:
1002:
923:
582:are a degree of structural organization of the
1586:
1275:
1273:
777:
714:promoter hypermethylation of tumor suppressor
675:digestion and next-generation DNA sequencing.
375:Sequence-Dependent Allele-Specific Methylation
354:and sciATAC-seq, scCOOL-seq, scNOMe-seq and sc
126:The main types of epigenetic changes include:
2129:
2086:
1388:
2475:
2331:"BioNews - Human Epigenome project launched"
1968:
1382:
999:
807:– Chromatin Immunoprecipitation Sequencing (
144:bases. This modification generally leads to
77:, is maintained through cell division (both
2089:"Emerging patterns of epigenomic variation"
1270:
964:
543:
1395:Microbiology and Molecular Biology Reviews
624:Thirdly, transcription start sites (TSS),
2507:
2458:
2409:
2399:
2381:
2307:
2258:
2217:
2169:
2136:Aguilar CA, Craighead HG (October 2013).
2112:
2038:
1916:
1867:
1818:
1769:
1720:
1671:
1622:
1612:
1563:
1503:
1414:
1365:
1305:
1253:
1170:
1160:
1114:
1104:
947:
289:
85:). The epigenome is essential for normal
2545:NCBI Gene Expression Omnibus Epigenomics
1587:Kim YJ, Cecchini KR, Kim TH (May 2011).
768:International Human Epigenome Consortium
681:
555:International Human Epigenome Consortium
55:transgenerational epigenetic inheritance
20:
1235:
411:
2594:
2240:
29:(blue) that can be methylated (green).
2555:Roadmap Epigenomics Visualization Hub
2426:
2362:
1890:
1525:
1523:
1438:
1436:
1434:
1327:
1325:
1208:10.1146/annurev.nutr.27.061406.093705
575:Topological associated domains (TADs)
310:level decreases sharply in the early
251:Factors affecting methylation pattern
2243:"Project set to map marks on genome"
1231:
1229:
1227:
1225:
1138:
1136:
1134:
1082:
1080:
1036:
1034:
182:
2281:
1389:Sterner DE, Berger SL (June 2000).
570:Chromatin interactions and domains;
61:and changes to the function of the
13:
1520:
1431:
1322:
129:
14:
2613:
2528:
1222:
1131:
1077:
1031:
1013:"Genetics, Epigenetic Mechanism"
901:Epigenome-wide association study
2375:
2343:
2324:
2275:
2234:
2186:
2055:
2006:
1933:
1884:
1835:
1786:
1737:
1688:
1639:
1580:
1479:
784:NIH Roadmap Epigenomics Project
694:is a currently active topic in
1897:Trends in Biochemical Sciences
1407:10.1128/MMBR.64.2.435-459.2000
1236:Wen L, Tang F (October 2019).
1187:
973:Journal of Cellular Physiology
702:undergo a major disruption of
580:Topological associated domains
518:
503:
332:CpG methylation is similar in
1:
2570:Epigenome Browser UCSC mirror
2087:Milosavljevic A (June 2011).
916:
757:Human Epigenome Pilot Project
734:drives aging by compromising
533:Non-coding RNA gene silencing
148:by preventing the binding of
2204:(7281): 587. February 2010.
1255:10.1016/j.molcel.2019.08.025
837:DNase I hypersensitive sites
751:As a prelude to a potential
651:specific regulatory events.
227:and quiescent regions (e.g.
7:
884:
778:Roadmap epigenomics project
746:
10:
2618:
2451:10.1016/j.cell.2012.12.033
2288:Genomics & Informatics
2241:Abbott A (February 2010).
2031:10.1016/j.cell.2022.12.027
1909:10.1016/j.tibs.2018.03.006
1811:10.1016/j.cell.2011.11.058
1664:10.1016/j.stem.2010.03.018
1505:10.1016/j.cell.2012.01.010
1196:Annual Review of Nutrition
949:10.1016/j.cell.2007.01.033
522:
507:
415:
133:
119:
2550:The Human Epigenome Atlas
2105:10.1016/j.tig.2011.03.001
2063:"Human Epigenome Project"
1992:10.1016/j.jid.2020.10.006
1860:10.1038/s41588-018-0278-6
1162:10.1186/s13059-019-1708-1
1106:10.1186/s13073-015-0209-4
932:"The mammalian epigenome"
686:
336:(GV) stage, intermediate
2401:10.3389/fgene.2014.00024
2382:Kanai Y, Arai E (2014).
2194:"Time for the epigenome"
1445:Nature Reviews. Genetics
1043:Nature Reviews. Genetics
725:
544:Structural modifications
453:Transcribed Gene Bodies
223:CoRSIVs are enriched in
115:
91:cellular differentiation
2582:Human Epigenome Project
1701:Genes & Development
1614:10.1073/pnas.1018279108
1298:10.1126/science.aar3146
1292:(6409). New York, N.Y.
832:Chromatin Accessibility
753:Human Epigenome Project
596:transcriptional control
390:gene regulatory circuit
2300:10.5808/GI.2013.11.1.7
2162:10.1038/nnano.2013.195
1891:Eagen KP (June 2018).
1762:10.4161/cc.10.16.17092
602:Firstly, they contain
340:(MI) stage and mature
290:Methylation in embryos
30:
2388:Frontiers in Genetics
2282:Bae JB (March 2013).
2142:Nature Nanotechnology
804:Histone Modifications
796:Human Epigenome Atlas
682:Clinical significance
669:chemical crosslinking
161:transposable elements
150:transcription factors
110:transposable elements
24:
2540:NCBI Epigenomics Hub
2488:Nature Biotechnology
1457:10.1038/nrg.2016.138
863:Small RNA Expression
418:Histone modification
412:Histone modification
382:bisulfite sequencing
75:histone modification
2210:2010Natur.463Q.587.
2154:2013NatNa...8..709A
1713:10.1101/gad.2005511
1605:2011PNAS..108.7391K
1556:10.1038/nature11082
1548:2012Natur.485..376D
1358:10.1038/nature14217
1350:2015Natur.518..350L
1055:10.1038/nrg.2016.45
879:Alzheimer's disease
525:Histone methylation
510:Histone acetylation
294:It was revealed by
2575:2021-02-14 at the
2355:2015-07-08 at the
2336:2010-12-28 at the
2093:Trends in Genetics
2025:(2): 305–326.e27.
789:2021-04-08 at the
673:restriction enzyme
671:of chromatin with
626:housekeeping genes
395:The study made by
31:
2494:(10): 1097–1105.
1980:J Invest Dermatol
1756:(16): 2779–2791.
1599:(18): 7391–7396.
1542:(7398): 376–380.
1344:(7539): 350–354.
985:10.1002/jcp.21678
896:Epigenome editing
857:expression arrays
501:
500:
456:Silenced Regions
450:Active Enhancers
447:Active Promoters
183:Methylation sites
2609:
2522:
2521:
2511:
2500:10.1038/nbt.1682
2479:
2473:
2472:
2462:
2430:
2424:
2423:
2413:
2403:
2379:
2373:
2366:
2360:
2347:
2341:
2328:
2322:
2321:
2311:
2279:
2273:
2272:
2262:
2238:
2232:
2231:
2221:
2190:
2184:
2183:
2173:
2133:
2127:
2126:
2116:
2084:
2078:
2077:
2075:
2074:
2065:. Archived from
2059:
2053:
2052:
2042:
2010:
2004:
2003:
1975:
1966:
1965:
1937:
1931:
1930:
1920:
1888:
1882:
1881:
1871:
1839:
1833:
1832:
1822:
1805:(1–2): 335–348.
1790:
1784:
1783:
1773:
1741:
1735:
1734:
1724:
1692:
1686:
1685:
1675:
1643:
1637:
1636:
1626:
1616:
1584:
1578:
1577:
1567:
1527:
1518:
1517:
1507:
1483:
1477:
1476:
1440:
1429:
1428:
1418:
1386:
1380:
1379:
1369:
1329:
1320:
1319:
1309:
1277:
1268:
1267:
1257:
1233:
1220:
1219:
1191:
1185:
1184:
1174:
1164:
1140:
1129:
1128:
1118:
1108:
1084:
1075:
1074:
1038:
1029:
1028:
1008:
997:
996:
968:
962:
961:
951:
927:
911:NCBI Epigenomics
782:One goal of the
698:research. Human
645:retrotransposons
444:
443:
334:germinal vesicle
235:regions, active
165:repeat sequences
157:5-methylcytosine
2617:
2616:
2612:
2611:
2610:
2608:
2607:
2606:
2592:
2591:
2587:Cancer Research
2577:Wayback Machine
2531:
2526:
2525:
2480:
2476:
2431:
2427:
2380:
2376:
2367:
2363:
2359:. 5 March 2010.
2357:Wayback Machine
2348:
2344:
2338:Wayback Machine
2329:
2325:
2280:
2276:
2260:10.1038/463596b
2253:(7281): 596–7.
2239:
2235:
2219:10.1038/463587a
2192:
2191:
2187:
2148:(10): 709–718.
2134:
2130:
2085:
2081:
2072:
2070:
2061:
2060:
2056:
2011:
2007:
1986:(4S): 961–967.
1976:
1969:
1938:
1934:
1889:
1885:
1848:Nature Genetics
1840:
1836:
1791:
1787:
1742:
1738:
1693:
1689:
1644:
1640:
1585:
1581:
1528:
1521:
1484:
1480:
1441:
1432:
1387:
1383:
1330:
1323:
1278:
1271:
1234:
1223:
1192:
1188:
1141:
1132:
1093:Genome Medicine
1085:
1078:
1039:
1032:
1009:
1000:
969:
965:
928:
924:
919:
906:Human epigenome
887:
848:Gene Expression
819:– Whole Genome
816:DNA Methylation
791:Wayback Machine
780:
749:
740:DNA replication
728:
704:DNA methylation
689:
684:
657:
577:
546:
535:
527:
521:
512:
506:
428:phosphorylation
420:
414:
377:
308:DNA methylation
302:process. After
292:
284:gene regulation
271:
253:
233:heterochromatic
203:nterindividual
185:
138:
136:DNA methylation
132:
130:DNA methylation
124:
118:
71:DNA methylation
67:human epigenome
17:
16:Biological term
12:
11:
5:
2615:
2605:
2604:
2590:
2589:
2584:
2579:
2567:
2562:
2557:
2552:
2547:
2542:
2537:
2530:
2529:External links
2527:
2524:
2523:
2474:
2445:(3): 642–654.
2425:
2374:
2361:
2342:
2323:
2274:
2233:
2185:
2128:
2099:(6): 242–250.
2079:
2054:
2005:
1967:
1954:10.1086/412317
1948:(3): 279–303.
1932:
1903:(6): 469–478.
1883:
1854:(1): 128–137.
1834:
1785:
1736:
1707:(7): 742–754.
1687:
1658:(5): 479–491.
1652:Cell Stem Cell
1638:
1579:
1519:
1498:(3): 458–472.
1478:
1430:
1381:
1321:
1269:
1248:(2): 320–328.
1242:Molecular Cell
1221:
1202:(1): 363–388.
1186:
1149:Genome Biology
1130:
1076:
1049:(6): 319–332.
1030:
998:
963:
942:(4): 669–681.
921:
920:
918:
915:
914:
913:
908:
903:
898:
893:
886:
883:
875:
874:
860:
844:
828:
812:
779:
776:
748:
745:
730:The idea that
727:
724:
688:
685:
683:
680:
656:
653:
608:RNA polymerase
576:
573:
572:
571:
568:
545:
542:
538:Non-coding RNA
534:
531:
523:Main article:
520:
517:
508:Main article:
505:
502:
499:
498:
493:
491:
486:
480:
479:
474:
469:
464:
458:
457:
454:
451:
448:
432:ubiquitination
416:Main article:
413:
410:
397:Onuchic et al.
376:
373:
296:immunostaining
291:
288:
270:
267:
252:
249:
184:
181:
146:gene silencing
134:Main article:
131:
128:
120:Main article:
117:
114:
15:
9:
6:
4:
3:
2:
2614:
2603:
2600:
2599:
2597:
2588:
2585:
2583:
2580:
2578:
2574:
2571:
2568:
2566:
2563:
2561:
2558:
2556:
2553:
2551:
2548:
2546:
2543:
2541:
2538:
2536:
2533:
2532:
2519:
2515:
2510:
2505:
2501:
2497:
2493:
2489:
2485:
2478:
2470:
2466:
2461:
2456:
2452:
2448:
2444:
2440:
2436:
2429:
2421:
2417:
2412:
2407:
2402:
2397:
2393:
2389:
2385:
2378:
2371:
2368:Eurice GmbH.
2365:
2358:
2354:
2351:
2346:
2339:
2335:
2332:
2327:
2319:
2315:
2310:
2305:
2301:
2297:
2293:
2289:
2285:
2278:
2270:
2266:
2261:
2256:
2252:
2248:
2244:
2237:
2229:
2225:
2220:
2215:
2211:
2207:
2203:
2199:
2195:
2189:
2181:
2177:
2172:
2167:
2163:
2159:
2155:
2151:
2147:
2143:
2139:
2132:
2124:
2120:
2115:
2110:
2106:
2102:
2098:
2094:
2090:
2083:
2069:on 2011-07-16
2068:
2064:
2058:
2050:
2046:
2041:
2036:
2032:
2028:
2024:
2020:
2016:
2009:
2001:
1997:
1993:
1989:
1985:
1981:
1974:
1972:
1963:
1959:
1955:
1951:
1947:
1943:
1936:
1928:
1924:
1919:
1914:
1910:
1906:
1902:
1898:
1894:
1887:
1879:
1875:
1870:
1865:
1861:
1857:
1853:
1849:
1845:
1838:
1830:
1826:
1821:
1816:
1812:
1808:
1804:
1800:
1796:
1789:
1781:
1777:
1772:
1767:
1763:
1759:
1755:
1751:
1747:
1740:
1732:
1728:
1723:
1718:
1714:
1710:
1706:
1702:
1698:
1691:
1683:
1679:
1674:
1669:
1665:
1661:
1657:
1653:
1649:
1642:
1634:
1630:
1625:
1620:
1615:
1610:
1606:
1602:
1598:
1594:
1590:
1583:
1575:
1571:
1566:
1561:
1557:
1553:
1549:
1545:
1541:
1537:
1533:
1526:
1524:
1515:
1511:
1506:
1501:
1497:
1493:
1489:
1482:
1474:
1470:
1466:
1462:
1458:
1454:
1450:
1446:
1439:
1437:
1435:
1426:
1422:
1417:
1412:
1408:
1404:
1401:(2): 435–59.
1400:
1396:
1392:
1385:
1377:
1373:
1368:
1363:
1359:
1355:
1351:
1347:
1343:
1339:
1335:
1328:
1326:
1317:
1313:
1308:
1303:
1299:
1295:
1291:
1287:
1283:
1276:
1274:
1265:
1261:
1256:
1251:
1247:
1243:
1239:
1232:
1230:
1228:
1226:
1217:
1213:
1209:
1205:
1201:
1197:
1190:
1182:
1178:
1173:
1168:
1163:
1158:
1154:
1150:
1146:
1139:
1137:
1135:
1126:
1122:
1117:
1112:
1107:
1102:
1098:
1094:
1090:
1083:
1081:
1072:
1068:
1064:
1060:
1056:
1052:
1048:
1044:
1037:
1035:
1026:
1022:
1018:
1014:
1007:
1005:
1003:
994:
990:
986:
982:
979:(2): 243–50.
978:
974:
967:
959:
955:
950:
945:
941:
937:
933:
926:
922:
912:
909:
907:
904:
902:
899:
897:
894:
892:
889:
888:
882:
880:
872:
868:
864:
861:
858:
854:
850:
849:
845:
842:
838:
834:
833:
829:
826:
822:
821:Bisulfite-Seq
818:
817:
813:
810:
806:
805:
801:
800:
799:
797:
792:
788:
785:
775:
773:
769:
764:
762:
758:
754:
744:
741:
737:
736:transcription
733:
723:
721:
718:, an altered
717:
713:
709:
705:
701:
697:
693:
679:
676:
674:
670:
665:
663:
652:
648:
646:
643:
639:
634:
631:
627:
622:
620:
615:
613:
609:
605:
600:
597:
592:
590:
585:
581:
569:
566:
563:
562:
561:
558:
556:
552:
541:
539:
530:
526:
516:
511:
497:
494:
492:
490:
487:
485:
482:
481:
478:
475:
473:
470:
468:
465:
463:
460:
459:
455:
452:
449:
446:
445:
442:
439:
437:
433:
429:
425:
419:
409:
405:
401:
398:
393:
391:
385:
383:
372:
370:
366:
361:
357:
353:
349:
345:
343:
339:
335:
330:
328:
324:
319:
317:
313:
309:
305:
304:fertilisation
301:
300:demethylation
297:
287:
285:
279:
277:
276:transcriptome
266:
263:
259:
258:polymorphisms
248:
244:
242:
238:
234:
230:
226:
221:
219:
215:
209:
206:
202:
198:
194:
190:
180:
178:
173:
168:
166:
162:
158:
153:
151:
147:
143:
137:
127:
123:
113:
111:
106:
104:
100:
96:
92:
88:
84:
80:
76:
72:
68:
64:
60:
56:
52:
48:
44:
40:
36:
28:
23:
19:
2491:
2487:
2477:
2442:
2438:
2428:
2391:
2387:
2377:
2370:"About IHEC"
2364:
2345:
2326:
2291:
2287:
2277:
2250:
2246:
2236:
2201:
2197:
2188:
2145:
2141:
2131:
2096:
2092:
2082:
2071:. Retrieved
2067:the original
2057:
2022:
2018:
2008:
1983:
1979:
1945:
1941:
1935:
1900:
1896:
1886:
1851:
1847:
1837:
1802:
1798:
1788:
1753:
1749:
1739:
1704:
1700:
1690:
1655:
1651:
1641:
1596:
1592:
1582:
1539:
1535:
1495:
1491:
1481:
1451:(1): 51–66.
1448:
1444:
1398:
1394:
1384:
1341:
1337:
1289:
1285:
1245:
1241:
1199:
1195:
1189:
1152:
1148:
1096:
1092:
1046:
1042:
1016:
976:
972:
966:
939:
935:
925:
876:
862:
846:
839:Sequencing (
830:
814:
802:
781:
765:
756:
752:
750:
729:
690:
677:
666:
662:interactomes
658:
649:
635:
623:
616:
601:
593:
578:
559:
547:
536:
528:
513:
440:
421:
406:
402:
396:
394:
386:
378:
346:
342:metaphase II
331:
320:
315:
293:
280:
272:
262:trans-acting
254:
247:cell types.
245:
229:subtelomeric
222:
218:chromosome 6
210:
204:
200:
196:
192:
188:
187:CoRSIVs are
186:
169:
154:
139:
125:
107:
69:, including
66:
38:
32:
18:
2602:Epigenetics
2294:(1): 7–14.
1017:StatPearls
891:Epigenetics
692:Epigenetics
640:/B1 and B2
519:Methylation
504:Acetylation
436:sumoylation
424:acetylation
338:metaphase I
216:) locus on
122:Epigenetics
87:development
2073:2011-06-29
1942:Q Rev Biol
1750:Cell Cycle
1155:(1): 105.
917:References
772:cell types
732:DNA damage
712:CpG island
589:Drosophila
565:Nucleosome
365:protamines
225:intergenic
195:egions of
1099:(1): 88.
867:smRNA-Seq
841:DNase-Seq
825:MeDIP-Seq
604:insulator
356:DNase-seq
348:Chromatin
312:pronuclei
241:enhancers
237:promoters
191:rrelated
177:phenotype
59:chromatin
51:expressed
39:epigenome
2596:Category
2573:Archived
2518:20852635
2469:23333102
2420:24592273
2353:Archived
2334:Archived
2318:23613677
2269:20162836
2228:20130607
2180:24091454
2123:21507501
2049:36638792
2040:10166133
2000:33494932
1927:29685368
1878:30478436
1829:22244452
1780:21822054
1731:21460038
1682:20452322
1633:21502535
1574:22495300
1514:22265598
1465:27867193
1425:10839822
1376:25693566
1316:30139913
1264:31563431
1216:17465856
1181:31155008
1125:26391122
1063:27156976
1025:30422591
993:19127539
958:17320505
885:See also
809:ChIP-Seq
787:Archived
747:Research
477:H3K27me3
472:H3K36me3
369:histones
352:ATAC-seq
199:ystemic
142:cytosine
27:histones
2509:2955169
2460:3563935
2411:3924033
2309:3630389
2206:Bibcode
2171:4072028
2150:Bibcode
2114:3104125
1962:7031747
1918:6028237
1869:6330062
1820:3368268
1771:3219543
1722:3070936
1673:2867844
1624:3088595
1601:Bibcode
1565:3356448
1544:Bibcode
1473:4461801
1367:4449149
1346:Bibcode
1307:6198826
1286:Science
1172:6545702
1116:4578259
853:RNA-Seq
720:histone
708:histone
619:H3K9me3
496:H3K9me3
489:H3K27ac
484:H3K27ac
467:H3K4me1
462:H3K4me3
325:and in
323:oocytes
316:de novo
83:meiosis
79:mitosis
47:histone
35:biology
2516:
2506:
2467:
2457:
2418:
2408:
2394:: 24.
2316:
2306:
2267:
2247:Nature
2226:
2198:Nature
2178:
2168:
2121:
2111:
2047:
2037:
1998:
1960:
1925:
1915:
1876:
1866:
1827:
1817:
1778:
1768:
1729:
1719:
1680:
1670:
1631:
1621:
1572:
1562:
1536:Nature
1512:
1471:
1463:
1423:
1413:
1374:
1364:
1338:Nature
1314:
1304:
1262:
1214:
1179:
1169:
1123:
1113:
1071:336906
1069:
1061:
1023:
991:
956:
871:miRNAs
761:genome
755:, the
700:tumors
696:cancer
687:Cancer
584:genome
551:ENCODE
434:, and
360:zygote
306:, the
239:, and
103:toxins
101:, and
99:stress
65:. The
63:genome
37:, the
1469:S2CID
1416:98999
1067:S2CID
726:Aging
716:genes
327:sperm
116:Types
2514:PMID
2465:PMID
2439:Cell
2416:PMID
2314:PMID
2265:PMID
2224:PMID
2176:PMID
2119:PMID
2045:PMID
2019:Cell
1996:PMID
1958:PMID
1923:PMID
1874:PMID
1825:PMID
1799:Cell
1776:PMID
1727:PMID
1678:PMID
1629:PMID
1570:PMID
1510:PMID
1492:Cell
1461:PMID
1421:PMID
1372:PMID
1312:PMID
1260:PMID
1212:PMID
1177:PMID
1121:PMID
1059:PMID
1021:PMID
989:PMID
954:PMID
936:Cell
855:and
738:and
706:and
642:SINE
630:tRNA
628:and
612:CTCF
163:and
95:diet
89:and
81:and
73:and
45:and
2504:PMC
2496:doi
2455:PMC
2447:doi
2443:152
2406:PMC
2396:doi
2304:PMC
2296:doi
2255:doi
2251:463
2214:doi
2202:463
2166:PMC
2158:doi
2109:PMC
2101:doi
2035:PMC
2027:doi
2023:186
1988:doi
1984:141
1950:doi
1913:PMC
1905:doi
1864:PMC
1856:doi
1815:PMC
1807:doi
1803:148
1766:PMC
1758:doi
1717:PMC
1709:doi
1668:PMC
1660:doi
1619:PMC
1609:doi
1597:108
1560:PMC
1552:doi
1540:485
1500:doi
1496:148
1453:doi
1411:PMC
1403:doi
1362:PMC
1354:doi
1342:518
1302:PMC
1294:doi
1290:361
1250:doi
1204:doi
1167:PMC
1157:doi
1111:PMC
1101:doi
1051:doi
981:doi
977:219
944:doi
940:128
835:–
638:Alu
367:by
214:MHC
172:CpG
43:DNA
33:In
2598::
2512:.
2502:.
2492:28
2490:.
2486:.
2463:.
2453:.
2441:.
2437:.
2414:.
2404:.
2390:.
2386:.
2312:.
2302:.
2292:11
2290:.
2286:.
2263:.
2249:.
2245:.
2222:.
2212:.
2200:.
2196:.
2174:.
2164:.
2156:.
2144:.
2140:.
2117:.
2107:.
2097:27
2095:.
2091:.
2043:.
2033:.
2021:.
2017:.
1994:.
1982:.
1970:^
1956:.
1946:56
1944:.
1921:.
1911:.
1901:43
1899:.
1895:.
1872:.
1862:.
1852:51
1850:.
1846:.
1823:.
1813:.
1801:.
1797:.
1774:.
1764:.
1754:10
1752:.
1748:.
1725:.
1715:.
1705:25
1703:.
1699:.
1676:.
1666:.
1654:.
1650:.
1627:.
1617:.
1607:.
1595:.
1591:.
1568:.
1558:.
1550:.
1538:.
1534:.
1522:^
1508:.
1494:.
1490:.
1467:.
1459:.
1449:18
1447:.
1433:^
1419:.
1409:.
1399:64
1397:.
1393:.
1370:.
1360:.
1352:.
1340:.
1336:.
1324:^
1310:.
1300:.
1288:.
1284:.
1272:^
1258:.
1246:76
1244:.
1240:.
1224:^
1210:.
1200:27
1198:.
1175:.
1165:.
1153:20
1151:.
1147:.
1133:^
1119:.
1109:.
1095:.
1091:.
1079:^
1065:.
1057:.
1047:17
1045:.
1033:^
1015:.
1001:^
987:.
975:.
952:.
938:.
934:.
881:.
865:–
851:–
774:.
614:.
591:.
430:,
426:,
371:.
286:.
189:Co
167:.
105:.
97:,
2520:.
2498::
2471:.
2449::
2422:.
2398::
2392:5
2372:.
2340:.
2320:.
2298::
2271:.
2257::
2230:.
2216::
2208::
2182:.
2160::
2152::
2146:8
2125:.
2103::
2076:.
2051:.
2029::
2002:.
1990::
1964:.
1952::
1929:.
1907::
1880:.
1858::
1831:.
1809::
1782:.
1760::
1733:.
1711::
1684:.
1662::
1656:6
1635:.
1611::
1603::
1576:.
1554::
1546::
1516:.
1502::
1475:.
1455::
1427:.
1405::
1378:.
1356::
1348::
1318:.
1296::
1266:.
1252::
1218:.
1206::
1183:.
1159::
1127:.
1103::
1097:7
1073:.
1053::
1027:.
995:.
983::
960:.
946::
873:.
212:(
205:V
201:I
197:S
193:R
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.