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Allis CD, Winston F, Berger SL, Workman JL (1997) Yeast Gcn5 functions in two
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1101/gad.11.13.1640
124. Vermeulen M, Eberl HC, Matarese F, Marks H, Denissov S, Butter F, Lee KK, Olsen JV,
Hyman AA, Stunnenberg HG, Mann M (2010) Quantitative interaction proteomics and
genome-wide profiling of epigenetic histone marks and their readers. Cell 142(6):967–980.
https://doi.org/10.1016/j.cell.2010.08.020
125. Law JA, Vashisht AA, Wohlschlegel JA, Jacobsen SE (2011) SHH1, a homeodomain protein
required for DNA methylation, as well as RDR2, RDM4, and chromatin remodeling factors,
associate with RNA polymerase IV. PLoS Genet 7(7):e1002195. https://doi.org/10.1371/
journal.pgen.1002195
126. Haag JR, Pikaard CS (2011) Multisubunit RNA polymerases IV and V: purveyors of
non-coding RNA for plant gene silencing. Nat Rev Mol Cell Biol 12(8):483–492. https://
doi.org/10.1038/nrm3152
127. Law JA, Du J, Hale CJ, Feng S, Krajewski K, Palanca AM, Strahl BD, Patel DJ, Jacobsen SE
(2013) Polymerase IV occupancy at RNA-directed DNA methylation sites requires SHH1.
Nature 498(7454):385–389. https://doi.org/10.1038/nature12178
128. Bostick M, Kim JK, Esteve PO, Clark A, Pradhan S, Jacobsen SE (2007) UHRF1 plays a role
in maintaining DNA methylation in mammalian cells. Science 317(5845):1760–1764. https://
doi.org/10.1126/science.1147939
129. Sharif J, Muto M, Takebayashi S, Suetake I, Iwamatsu A, Endo TA, Shinga J,
Mizutani-Koseki Y, Toyoda T, Okamura K, Tajima S, Mitsuya K, Okano M, Koseki H
(2007) The SRA protein Np95 mediates epigenetic inheritance by recruiting Dnmt1 to
methylated DNA. Nature 450(7171):908–912. https://doi.org/10.1038/nature06397
130. Arita K, Isogai S, Oda T, Unoki M, Sugita K, Sekiyama N, Kuwata K, Hamamoto R,
Tochio H, Sato M, Ariyoshi M, Shirakawa M (2012) Recognition of modification status
on a histone H3 tail by linked histone reader modules of the epigenetic regulator UHRF1.
Proc Natl Acad Sci U S A 109(32):12950–12955. https://doi.org/10.1073/pnas.1203701109
131. Berkyurek AC, Suetake I, Arita K, Takeshita K, Nakagawa A, Shirakawa M, Tajima S (2014)
The DNA methyltransferase Dnmt1 directly interacts with the SET and RING fingerassociated (SRA) domain of the multifunctional protein Uhrf1 to facilitate accession of
the catalytic center to hemi-methylated DNA. J Biol Chem 289(1):379–386. https://doi.org/
10.1074/jbc.M113.523209
132. Bashtrykov P, Jankevicius G, Jurkowska RZ, Ragozin S, Jeltsch A (2014) The UHRF1 protein
stimulates the activity and specificity of the maintenance DNA methyltransferase DNMT1
by an allosteric mechanism. J Biol Chem 289(7):4106–4115. https://doi.org/10.1074/jbc.
M113.528893
133. Nady N, Lemak A, Walker JR, Avvakumov GV, Kareta MS, Achour M, Xue S, Duan S,
Allali-Hassani A, Zuo X, Wang YX, Bronner C, Chedin F, Arrowsmith CH, Dhe-Paganon S
(2011) Recognition of multivalent histone states associated with heterochromatin by UHRF1
protein. J Biol Chem 286(27):24300–24311. https://doi.org/10.1074/jbc.M111.234104
134. Nishiyama A, Yamaguchi L, Sharif J, Johmura Y, Kawamura T, Nakanishi K, Shimamura S,
Arita K, Kodama T, Ishikawa F, Koseki H, Nakanishi M (2013) Uhrf1-dependent
H3K23 ubiquitylation couples maintenance DNA methylation and replication. Nature
502(7470):249–253. https://doi.org/10.1038/nature12488
135. Liu X, Gao Q, Li P, Zhao Q, Zhang J, Li J, Koseki H, Wong J (2013) UHRF1 targets DNMT1
for DNA methylation through cooperative binding of hemi-methylated DNA and methylated
H3K9. Nat Commun 4:1563. https://doi.org/10.1038/ncomms2562
136. Lallous N, Legrand P, McEwen AG, Ramon-Maiques S, Samama JP, Birck C (2011) The
PHD finger of human UHRF1 reveals a new subgroup of unmethylated histone H3 tail readers.
PLoS One 6(11):e27599. https://doi.org/10.1371/journal.pone.0027599
Methyl-Readers and Inhibitors
391
Allis CD, Winston F, Berger SL, Workman JL (1997) Yeast Gcn5 functions in two
multisubunit complexes to acetylate nucleosomal histones: characterization of an Ada complex and the SAGA (Spt/Ada) complex. Genes Dev 11(13):1640–1650. https://doi.org/10.
1101/gad.11.13.1640
124. Vermeulen M, Eberl HC, Matarese F, Marks H, Denissov S, Butter F, Lee KK, Olsen JV,
Hyman AA, Stunnenberg HG, Mann M (2010) Quantitative interaction proteomics and
genome-wide profiling of epigenetic histone marks and their readers. Cell 142(6):967–980.
https://doi.org/10.1016/j.cell.2010.08.020
125. Law JA, Vashisht AA, Wohlschlegel JA, Jacobsen SE (2011) SHH1, a homeodomain protein
required for DNA methylation, as well as RDR2, RDM4, and chromatin remodeling factors,
associate with RNA polymerase IV. PLoS Genet 7(7):e1002195. https://doi.org/10.1371/
journal.pgen.1002195
126. Haag JR, Pikaard CS (2011) Multisubunit RNA polymerases IV and V: purveyors of
non-coding RNA for plant gene silencing. Nat Rev Mol Cell Biol 12(8):483–492. https://
doi.org/10.1038/nrm3152
127. Law JA, Du J, Hale CJ, Feng S, Krajewski K, Palanca AM, Strahl BD, Patel DJ, Jacobsen SE
(2013) Polymerase IV occupancy at RNA-directed DNA methylation sites requires SHH1.
Nature 498(7454):385–389. https://doi.org/10.1038/nature12178
128. Bostick M, Kim JK, Esteve PO, Clark A, Pradhan S, Jacobsen SE (2007) UHRF1 plays a role
in maintaining DNA methylation in mammalian cells. Science 317(5845):1760–1764. https://
doi.org/10.1126/science.1147939
129. Sharif J, Muto M, Takebayashi S, Suetake I, Iwamatsu A, Endo TA, Shinga J,
Mizutani-Koseki Y, Toyoda T, Okamura K, Tajima S, Mitsuya K, Okano M, Koseki H
(2007) The SRA protein Np95 mediates epigenetic inheritance by recruiting Dnmt1 to
methylated DNA. Nature 450(7171):908–912. https://doi.org/10.1038/nature06397
130. Arita K, Isogai S, Oda T, Unoki M, Sugita K, Sekiyama N, Kuwata K, Hamamoto R,
Tochio H, Sato M, Ariyoshi M, Shirakawa M (2012) Recognition of modification status
on a histone H3 tail by linked histone reader modules of the epigenetic regulator UHRF1.
Proc Natl Acad Sci U S A 109(32):12950–12955. https://doi.org/10.1073/pnas.1203701109
131. Berkyurek AC, Suetake I, Arita K, Takeshita K, Nakagawa A, Shirakawa M, Tajima S (2014)
The DNA methyltransferase Dnmt1 directly interacts with the SET and RING fingerassociated (SRA) domain of the multifunctional protein Uhrf1 to facilitate accession of
the catalytic center to hemi-methylated DNA. J Biol Chem 289(1):379–386. https://doi.org/
10.1074/jbc.M113.523209
132. Bashtrykov P, Jankevicius G, Jurkowska RZ, Ragozin S, Jeltsch A (2014) The UHRF1 protein
stimulates the activity and specificity of the maintenance DNA methyltransferase DNMT1
by an allosteric mechanism. J Biol Chem 289(7):4106–4115. https://doi.org/10.1074/jbc.
M113.528893
133. Nady N, Lemak A, Walker JR, Avvakumov GV, Kareta MS, Achour M, Xue S, Duan S,
Allali-Hassani A, Zuo X, Wang YX, Bronner C, Chedin F, Arrowsmith CH, Dhe-Paganon S
(2011) Recognition of multivalent histone states associated with heterochromatin by UHRF1
protein. J Biol Chem 286(27):24300–24311. https://doi.org/10.1074/jbc.M111.234104
134. Nishiyama A, Yamaguchi L, Sharif J, Johmura Y, Kawamura T, Nakanishi K, Shimamura S,
Arita K, Kodama T, Ishikawa F, Koseki H, Nakanishi M (2013) Uhrf1-dependent
H3K23 ubiquitylation couples maintenance DNA methylation and replication. Nature
502(7470):249–253. https://doi.org/10.1038/nature12488
135. Liu X, Gao Q, Li P, Zhao Q, Zhang J, Li J, Koseki H, Wong J (2013) UHRF1 targets DNMT1
for DNA methylation through cooperative binding of hemi-methylated DNA and methylated
H3K9. Nat Commun 4:1563. https://doi.org/10.1038/ncomms2562
136. Lallous N, Legrand P, McEwen AG, Ramon-Maiques S, Samama JP, Birck C (2011) The
PHD finger of human UHRF1 reveals a new subgroup of unmethylated histone H3 tail readers.
PLoS One 6(11):e27599. https://doi.org/10.1371/journal.pone.0027599
Methyl-Readers and Inhibitors
391