Nucleosome destabilization by nuclear non-coding RNAs.

Cell Line Cell Nucleus / genetics Chromatin / chemistry Chromatin Assembly and Disassembly Genetic Loci Histones / metabolism Humans In Situ Hybridization, Fluorescence Nucleic Acid Conformation Nucleosomes / genetics Protein Stability RNA, Untranslated / chemistry

Journal

Communications biology
ISSN: 2399-3642
Titre abrégé: Commun Biol
Pays: England
ID NLM: 101719179

Informations de publication

Date de publication:
11 02 2020
Historique:
received: 03 06 2019
accepted: 16 01 2020
entrez: 13 2 2020
pubmed: 13 2 2020
medline: 11 5 2021
Statut: epublish

Résumé

In the nucleus, genomic DNA is wrapped around histone octamers to form nucleosomes. In principle, nucleosomes are substantial barriers to transcriptional activities. Nuclear non-coding RNAs (ncRNAs) are proposed to function in chromatin conformation modulation and transcriptional regulation. However, it remains unclear how ncRNAs affect the nucleosome structure. Eleanors are clusters of ncRNAs that accumulate around the estrogen receptor-α (ESR1) gene locus in long-term estrogen deprivation (LTED) breast cancer cells, and markedly enhance the transcription of the ESR1 gene. Here we detected nucleosome depletion around the transcription site of Eleanor2, the most highly expressed Eleanor in the LTED cells. We found that the purified Eleanor2 RNA fragment drastically destabilized the nucleosome in vitro. This activity was also exerted by other ncRNAs, but not by poly(U) RNA or DNA. The RNA-mediated nucleosome destabilization may be a common feature among natural nuclear RNAs, and may function in transcription regulation in chromatin.

Identifiants

DOI: 10.1038/s42003-020-0784-9 PMID: 32047236 PMC: PMC7012929
pubmed: 32047236
doi: 10.1038/s42003-020-0784-9
pii: 10.1038/s42003-020-0784-9
pmc: PMC7012929
doi:

Substances chimiques

Chromatin 0
Histones 0
Nucleosomes 0
RNA, Untranslated 0

Types de publication

Journal Article Research Support, Non-U.S. Gov't

Langues

eng

Sous-ensembles de citation

IM

Pagination

60

Références

Wolffe, A. P. Chromatin: Structure & Function (Academic Press, London, UK, 1998).
Luger, K., Mäder, A. W., Richmond, R. K., Sargent, D. F. & Richmond, T. J. Crystal structure of the nucleosome core particle at 2.8 Å resolution. Nature 389, 251–260 (1997).
doi: 10.1038/38444 pubmed: 9305837
Bondarenko, V. A. et al. Nucleosomes can form a polar barrier to transcript elongation by RNA polymerase II. Mol. Cell 24, 469–479 (2006).
pubmed: 17081995 doi: 10.1016/j.molcel.2006.09.009 pmcid: 17081995
Luger, K., Dechassa, M. L. & Tremethick, D. J. New insights into nucleosome and chromatin structure: an ordered state or a disordered affair? Nat. Rev. Mol. Cell Biol. 13, 436–447 (2012).
pubmed: 22722606 pmcid: 3408961 doi: 10.1038/nrm3382
Petesch, S. J. & Lis, J. T. Overcoming the nucleosome barrier during transcript elongation. Trends Genet. 28, 285–294 (2012).
pubmed: 22465610 pmcid: 3466053 doi: 10.1016/j.tig.2012.02.005
Wilusz, J. E., Sunwoo, H. & Spector, D. L. Long noncoding RNAs: functional surprises from the RNA world. Genes Dev. 23, 1494–1504 (2009).
pubmed: 19571179 pmcid: 3152381 doi: 10.1101/gad.1800909
Rinn, J. L. & Chang, H. Y. Genome regulation by long noncoding RNAs. Annu. Rev. Biochem. 81, 145–166 (2012).
pubmed: 22663078 doi: 10.1146/annurev-biochem-051410-092902 pmcid: 22663078
Ding, D. Q. et al. Meiosis-specific noncoding RNA mediates robust pairing of homologous chromosomes in meiosis. Science 11, 732–736 (2012).
doi: 10.1126/science.1219518
Batista, P. J. & Chang, H. Y. Long noncoding RNAs: cellular address codes in development and disease. Cell 152, 1298–1307 (2013).
pubmed: 3651923 pmcid: 3651923 doi: 10.1016/j.cell.2013.02.012
Ulitsky, I. & Bartel, D. P. lincRNAs: genomics, evolution, and mechanisms. Cell 154, 26–46 (2013).
pubmed: 3924787 pmcid: 3924787 doi: 10.1016/j.cell.2013.06.020
Cech, T. R. & Steitz, J. A. The noncoding RNA revolution-trashing old rules to forge new ones. Cell 157, 77–94 (2014).
pubmed: 24679528 doi: 10.1016/j.cell.2014.03.008
Krawczyk, M. & Emerson, B. M. p50-associated COX-2 extragenic RNA (PACER) activates COX-2 gene expression by occluding repressive NF-kappaB complexes. eLife 3, e01776 (2014).
pubmed: 24843008 pmcid: 4017649 doi: 10.7554/eLife.01776
Kopp, F. & Mendell, J. T. Functional classification and experimental dissection of long noncoding RNAs. Cell 172, 393–407 (2018).
pubmed: 29373828 pmcid: 5978744 doi: 10.1016/j.cell.2018.01.011
The ENCODE Project Consortium. An integrated encyclopedia of DNA elements in the human genome. Nature 489, 57–74 (2012).
pmcid: 3439153 doi: 10.1038/nature11247 pubmed: 3439153
Djebali, S. et al. Landscape of transcription in human cells. Nature 489, 101–108 (2012).
pubmed: 3684276 pmcid: 3684276 doi: 10.1038/nature11233
Bánfai, B. et al. Long noncoding RNAs are rarely translated in two human cell lines. Genome Res. 22, 1646–1657 (2012).
pubmed: 22955977 pmcid: 3431482 doi: 10.1101/gr.134767.111
Carninci, P. et al. The transcriptional landscape of the mammalian genome. Science 309, 1559–1563 (2005).
pubmed: 16141072 doi: 10.1126/science.1112014 pmcid: 16141072
Derrien, T. et al. The GENCODE v7 catalog of human long noncoding RNAs: analysis of their gene structure, evolution, and expression. Genome Res. 22, 1775–1789 (2012).
pubmed: 22955988 pmcid: 3431493 doi: 10.1101/gr.132159.111
Jeng, M. H. et al. Estrogen receptor expression and function in long-term estrogen-deprived human breast cancer cells. Endocrinology 139, 4164–4174 (1998).
pubmed: 9751496 doi: 10.1210/endo.139.10.6229 pmcid: 9751496
Chan, C. M., Martin, L. A., Johnston, S. R., Ali, S. & Dowsett, M. Molecular changes associated with the acquisition of oestrogen hypersensitivity in MCF-7 breast cancer cells on long-term oestrogen deprivation. J. Steroid Biochem. Mol. Biol. 81, 333–341 (2002).
pubmed: 12361723 doi: 10.1016/S0960-0760(02)00074-2
Martin, L. A. et al. Enhanced estrogen receptor (ER) alpha, ERBB2, and MAPK signal transduction pathways operate during the adaptation of MCF-7 cells to long term estrogen deprivation. J. Biol. Chem. 278, 30458–30468 (2003).
pubmed: 12775708 doi: 10.1074/jbc.M305226200
Martin, L. A., Farmer, I., Johnston, S. R., Ali, S. & Dowsett, M. Elevated ERK1/ERK2/estrogen receptor cross-talk enhances estrogen-mediated signaling during long-term estrogen deprivation. Endocr. Relat. Cancer 12, S75–S84 (2005).
pubmed: 16113101 doi: 10.1677/erc.1.01023
Aguilar, H. et al. Biological reprogramming in acquired resistance to endocrine therapy of breast cancer. Oncogene 29, 6071–6083 (2010).
pubmed: 20711236 doi: 10.1038/onc.2010.333
Magnani, L. et al. Genome-wide reprogramming of the chromatin landscape underlies endocrine therapy resistance in breast cancer. Proc. Natl Acad. Sci. USA 110, E1490–E1499 (2013).
pubmed: 23576735 doi: 10.1073/pnas.1219992110
Tomita, S. et al. A cluster of noncoding RNAs activates the ESR1 locus during breast cancer adaptation. Nat. Commun. 6, 6966 (2015).
pubmed: 25923108 pmcid: 4421845 doi: 10.1038/ncomms7966
West, J. A. et al. The long noncoding RNAs NEAT1 and MALAT1 bind active chromatin sites. Mol. Cell 55, 791–802 (2014).
pubmed: 25155612 pmcid: 4428586 doi: 10.1016/j.molcel.2014.07.012
Miyagawa, R. et al. Identification of cis- and trans-acting factors involved in the localization of MALAT-1 noncoding RNA to nuclear speckles. RNA 18, 738–751 (2012).
pubmed: 22355166 pmcid: 3312561 doi: 10.1261/rna.028639.111
Rhie, S. K. et al. Identification of activated enhancers and linked transcription factors in breast, prostate, and kidney tumors by tracing enhancer networks using epigenetic traits. Epigenetics Chromatin 9, 50 (2016).
pubmed: 27833659 pmcid: 5103450 doi: 10.1186/s13072-016-0102-4
Niknafs, Y. S. et al. The lncRNA landscape of breast cancer reveals a role for DSCAM-AS1 in breast cancer progression. Nat. Commun. 7, 12791 (2016).
pubmed: 27666543 pmcid: 5052669 doi: 10.1038/ncomms12791
Iyer, M. K. et al. The landscape of long noncoding RNAs in the human transcriptome. Nat. Genet. 47, 199–208 (2015).
pubmed: 25599403 pmcid: 4417758 doi: 10.1038/ng.3192
Dyer, P. N. et al. Reconstitution of nucleosome core particles from recombinant histones and DNA. Methods Enzymol. 375, 23–44 (2004).
pubmed: 14870657 doi: 10.1016/S0076-6879(03)75002-2 pmcid: 14870657
Taguchi, H., Horikoshi, N., Arimura, Y. & Kurumizaka, H. A method for evaluating nucleosome stability with a protein-binding fluorescent dye. Methods 70, 119–126 (2014).
pubmed: 25220913 doi: 10.1016/j.ymeth.2014.08.019 pmcid: 25220913
Frouws, T. D., Duda, S. C. & Richmond, T. J. X-ray structure of the MMTV-A nucleosome core. Proc. Natl Acad. Sci. USA 113, 1214–1219 (2016).
pubmed: 26787910 doi: 10.1073/pnas.1524607113 pmcid: 26787910
Ji, P. et al. MALAT-1, a novel noncoding RNA, and thymosin beta4 predict metastasis and survival in early-stage non-small cell lung cancer. Oncogene 22, 8031–8041 (2003).
pubmed: 12970751 doi: 10.1038/sj.onc.1206928 pmcid: 12970751
Yoshimoto, R., Mayeda, A., Yoshida, M. & Nakagawa, S. MALAT1 long non-coding RNA in cancer. Biochim. Biophys. Acta 1859, 192–199 (2016).
pubmed: 26434412 doi: 10.1016/j.bbagrm.2015.09.012 pmcid: 26434412
Sun, Q., Hao, Q. & Prasanth, K. V. Nuclear long noncoding RNAs: key regulators of gene expression. Trends Genet. 34, 142–157 (2017).
doi: 10.1016/j.tig.2017.11.005
Misteli, T. et al. The dynamics of a pre-mRNA splicing factor in living cells. Nature 387, 523–527 (1997).
pubmed: 9168118 doi: 10.1038/387523a0 pmcid: 9168118
Shopland, L. S. et al. Clustering of multiple specific genes and gene-rich R-bands around SC-35 domains: evidence for local euchromatic neighborhoods. J. Cell Biol. 162, 981–990 (2003).
pubmed: 12975345 pmcid: 2172856 doi: 10.1083/jcb.200303131
Wutz, A. Gene silencing in X-chromosome inactivation: advances in understanding facultative heterochromatin formation. Nat. Rev. Genet. 12, 542–553 (2011).
pubmed: 21765457 doi: 10.1038/nrg3035 pmcid: 21765457
Zhang, B. et al. PHGDH defines a metabolic subtype in lung adenocarcinomas with poor prognosis. Cell Rep. 19, 2289–2303 (2017).
pubmed: 28614715 doi: 10.1016/j.celrep.2017.05.067 pmcid: 28614715
Richards, E. G., Flessel, C. P. & Fresco, J. R. Polynucleotides. VI. Molecular properties and conformation of polyribouridylic acid. Biopolymers 1, 431–446 (1963).
doi: 10.1002/bip.360010504
Esteller, M. Non-coding RNAs in human disease. Nat. Rev. Genet. 12, 861–874 (2011).
pubmed: 22094949 doi: 10.1038/nrg3074 pmcid: 22094949
Abdalla, M. O. A. et al. The Eleanor ncRNAs activate the topological domain of the ESR1 locus to balance against apoptosis. Nat. Commun. 10, 3778 (2019).
pubmed: 31439835 pmcid: 6706407 doi: 10.1038/s41467-019-11378-4
Yamamoto, T. et al. Endocrine therapy-resistant breast cancer model cells are inhibited by soybean glyceollin I through Eleanor non-coding RNA. Sci. Rep. 8, 15202 (2018).
pubmed: 30315184 pmcid: 6185934 doi: 10.1038/s41598-018-33227-y
Li, X. et al. GRID-seq reveals the global RNA-chromatin interactome. Nat. Biotechnol. 35, 940–950 (2017).
pubmed: 28922346 pmcid: 5953555 doi: 10.1038/nbt.3968
Wang, K. A long noncoding RNA maintains active chromatin to coordinate homeotic gene expression. Nature 472, 120–124 (2011).
pubmed: 21423168 pmcid: 3670758 doi: 10.1038/nature09819
Li, W. et al. Functional roles of enhancer RNAs for oestrogen-dependent transcriptional activation. Nature 498, 516–520 (2013).
pubmed: 23728302 pmcid: 3718886 doi: 10.1038/nature12210
Davidovich, C., Zheng, L., Goodrich, K. J. & Cech, T. R. Promiscuous RNA binding by Polycomb repressive complex 2. Nat. Struct. Mol. Biol. 20, 1250–1257 (2013).
pubmed: 24077223 pmcid: 3823624 doi: 10.1038/nsmb.2679
Cifuentes-Rojas, C., Hernandez, A. J., Sarma, K. & Lee, J. T. Regulatory interactions between RNA and polycomb repressive complex 2. Mol. Cell 55, 171–185 (2014).
pubmed: 24882207 pmcid: 4107928 doi: 10.1016/j.molcel.2014.05.009
Holoch, D. & Margueron, R. Mechanisms regulating PRC2 recruitment and enzymatic activity. Trends Biochem. Sci. 42, 531–542 (2017).
pubmed: 28483375 pmcid: 28483375 doi: 10.1016/j.tibs.2017.04.003
Zhao, J., Sun, B. K., Erwin, J. A., Song, J. J. & Lee, J. T. Polycomb proteins targeted by a short repeat RNA to the mouse X chromosome. Science 322, 750–756 (2008).
pubmed: 18974356 pmcid: 2748911 doi: 10.1126/science.1163045
Pandey, R. R. et al. Kcnq1ot1 antisense noncoding RNA mediates lineage-specific transcriptional silencing through chromatin-level regulation. Mol. Cell 32, 232–246 (2008).
pubmed: 18951091 doi: 10.1016/j.molcel.2008.08.022
Yap, K. L. et al. Molecular interplay of the noncoding RNA ANRIL and methylated histone H3 lysine 27 by polycomb CBX7 in transcriptional silencing of INK4a. Mol. Cell 38, 662–674 (2010).
pubmed: 20541999 pmcid: 2886305 doi: 10.1016/j.molcel.2010.03.021
O’Leary, V. B. et al. PARTICLE, a triplex-forming long ncRNA, regulates locus-specific methylation in response to low-dose irradiation. Cell Rep. 11, 474–485 (2015).
pubmed: 25900080 doi: 10.1016/j.celrep.2015.03.043
Beltran, M. et al. The interaction of PRC2 with RNA or chromatin is mutually antagonistic. Genome Res. 26, 896–907 (2016).
pubmed: 27197219 pmcid: 4937559 doi: 10.1101/gr.197632.115
Han, P. & Chang, C. P. Long non-coding RNA and chromatin remodelling. RNA Biol. 12, 1094–1098 (2015).
pubmed: 26177256 pmcid: 4829272 doi: 10.1080/15476286.2015.1063770
Audia, J. E. & Campbell, R. M. Histone modifications and cancer. Cold Spring Harb. Perspect. Biol. 8, a019521 (2016).
pubmed: 27037415 pmcid: 4817802 doi: 10.1101/cshperspect.a019521
Simon, M. et al. Histone fold modifications control nucleosome unwrapping and disassembly. Proc. Natl Acad. Sci. USA 108, 12711–12716 (2011).
pubmed: 21768347 doi: 10.1073/pnas.1106264108
Krajewski, W. A., Li, J. & Dou, Y. Effects of histone H2B ubiquitylation on the nucleosome structure and dynamics. Nucleic Acids Res. 46, 7631–7642 (2018).
pubmed: 29931239 pmcid: 6125632 doi: 10.1093/nar/gky526
Li, H. & Durbin, R. Fast and accurate short read alignment with Burrows-Wheeler transform. Bioinformatics 25, 1754–1760 (2009).
pubmed: 19451168 pmcid: 19451168
Robinson, J. T. et al. Integrative genomics viewer. Nat. Biotechnol. 29, 24–26 (2011).
pubmed: 21221095 pmcid: 3346182 doi: 10.1038/nbt.1754
Thorvaldsdóttir, H., Robinson, J. T. & Mesirov, J. P. Integrative genomics viewer (IGV): high-performance genomics data visualization and exploration. Brief. Bioinform. 14, 178–192 (2013).
pubmed: 22517427 doi: 10.1093/bib/bbs017
Tanaka, Y. et al. Expression and purification of recombinant human histones. Methods 33, 3–11 (2004).
pubmed: 15039081 doi: 10.1016/j.ymeth.2003.10.024
Machida, S. et al. Structural basis of heterochromatin formation by human HP1. Mol. Cell 69, 385–397 (2018).
pubmed: 29336876 doi: 10.1016/j.molcel.2017.12.011
Tachiwana, H. et al. Structural basis of instability of the nucleosome containing a testis-specific histone variant, human H3T. Proc. Natl Acad. Sci. USA 107, 10454–10459 (2010).
pubmed: 20498094 doi: 10.1073/pnas.1003064107
Hamada, M., Kiryu, H., Sato, K., Mituyama, T. & Asai, K. Prediction of RNA secondary structure using generalized centroid estimators. Bioinformatics 25, 465–473 (2009).
pubmed: 19095700 doi: 10.1093/bioinformatics/btn601
McCaskill, J. S. The equilibrium partition function and base pair binding probabilities for RNA secondary structure. Biopolymers 29, 1105–1119 (1990).
pubmed: 1695107 doi: 10.1002/bip.360290621
Corces, M. R. et al. An improved ATAC-seq protocol reduces background and enables interrogation of frozen tissues. Nat. Methods 14, 959–962 (2017).
pubmed: 28846090 pmcid: 5623106 doi: 10.1038/nmeth.4396
Langmead, B. & Salzberg, S. L. Fast gapped-read alignment with Bowtie 2. Nat. Methods 9, 357–359 (2012).
pubmed: 3322381 pmcid: 3322381 doi: 10.1038/nmeth.1923
Ramírez, F., Dündar, F., Diehl, S., Grüning, B. A. & Manke, T. deepTools: a flexible platform for exploring deep-sequencing data. Nucleic Acids Res. 42, W187–191 (2014).
pubmed: 24799436 pmcid: 4086134 doi: 10.1093/nar/gku365

Auteurs

Risa Fujita (R)

Laboratory of Chromatin Structure and Function, Institute for Quantitative Biosciences, The University of Tokyo, 1-1-1 Yayoi, Bunkyo-ku, Tokyo, 113-0032, Japan.

Tatsuro Yamamoto (T)

Department of Medical Cell Biology, Institute of Molecular Embryology and Genetics, Kumamoto University, 2-2-1 Honjo, Chuo-ku, Kumamoto, 860-0811, Japan.
Division of Cancer Biology, The Cancer Institute of Japanese Foundation for Cancer Research, 3-8-31 Ariake, Koto-ku, Tokyo, 135-8550, Japan.

Yasuhiro Arimura (Y)

Laboratory of Chromatin Structure and Function, Institute for Quantitative Biosciences, The University of Tokyo, 1-1-1 Yayoi, Bunkyo-ku, Tokyo, 113-0032, Japan.

Saori Fujiwara (S)

Department of Medical Cell Biology, Institute of Molecular Embryology and Genetics, Kumamoto University, 2-2-1 Honjo, Chuo-ku, Kumamoto, 860-0811, Japan.

Hiroaki Tachiwana (H)

Division of Cancer Biology, The Cancer Institute of Japanese Foundation for Cancer Research, 3-8-31 Ariake, Koto-ku, Tokyo, 135-8550, Japan.

Yuichi Ichikawa (Y)

Division of Cancer Biology, The Cancer Institute of Japanese Foundation for Cancer Research, 3-8-31 Ariake, Koto-ku, Tokyo, 135-8550, Japan.

Yuka Sakata (Y)

Division of Cancer Biology, The Cancer Institute of Japanese Foundation for Cancer Research, 3-8-31 Ariake, Koto-ku, Tokyo, 135-8550, Japan.
ORCID: 0000-0003-4254-2417

Liying Yang (L)

Project for Cancer Epigenomics, The Cancer Institute of Japanese Foundation for Cancer Research, 3-8-31 Ariake, Koto-ku, Tokyo, 135-8550, Japan.

Reo Maruyama (R)

Project for Cancer Epigenomics, The Cancer Institute of Japanese Foundation for Cancer Research, 3-8-31 Ariake, Koto-ku, Tokyo, 135-8550, Japan.

Michiaki Hamada (M)

Graduate School of Advanced Science and Engineering, Waseda University, 3-4-1 Okubo, Shinjuku-ku, Tokyo, 169-8555, Japan.
AIST-Waseda University Computational Bio Big-Data Open Innovation Laboratory (CBBD-OIL), 3-4-1 Okubo, Shinjuku-ku, Tokyo, 169-8555, Japan.

Mitsuyoshi Nakao (M)

Department of Medical Cell Biology, Institute of Molecular Embryology and Genetics, Kumamoto University, 2-2-1 Honjo, Chuo-ku, Kumamoto, 860-0811, Japan.

Noriko Saitoh (N)

Division of Cancer Biology, The Cancer Institute of Japanese Foundation for Cancer Research, 3-8-31 Ariake, Koto-ku, Tokyo, 135-8550, Japan. noriko.saitoh@jfcr.or.jp.

Hitoshi Kurumizaka (H)

Laboratory of Chromatin Structure and Function, Institute for Quantitative Biosciences, The University of Tokyo, 1-1-1 Yayoi, Bunkyo-ku, Tokyo, 113-0032, Japan. kurumizaka@iam.u-tokyo.ac.jp.
ORCID: 0000-0001-7412-3722

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Classifications MeSH

questionsmedicales.fr › Cellules › Cellules cultivées › Lignée cellulaire › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Cellules › Structures cellulaires › Espace intracellulaire › Cytoplasme › Structures cytoplasmiques › Organites › Noyau de la cellule › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Phénomènes génétiques › Structures génétiques › Structures de chromosome › Chromatine › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Phénomènes génétiques › Régulation de l'expression des gènes › Assemblage et désassemblage de la chromatine › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Phénomènes génétiques › Structures génétiques › Génome › Composants de génome › Locus génétiques › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Acides aminés, peptides et protéines › Protéines › Nucléoprotéines › Histone › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Eucaryotes › Animaux › Chordés › Vertébrés › Mammifères › Eutheria › Primates › Haplorhini › Catarrhini › Hominidae › Humains › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Techniques d'investigation › Techniques génétiques › Hybridation in situ › Hybridation fluorescente in situ › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Phénomènes génétiques › Structures génétiques › Conformation d'acide nucléique › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Phénomènes génétiques › Structures génétiques › Structures de chromosome › Chromatine › Nucléosomes › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Phénomènes chimiques › Phénomènes biochimiques › Stabilité protéique › Nucleosome destabilization by nuclear non-coding RNAs.
questionsmedicales.fr › Acides nucléiques, nucléotides et nucléosides › Acides nucléiques › ARN › ARN non traduit › Nucleosome destabilization by nuclear non-coding RNAs.