Table 1.
List of mitotic histone PTMs (identified in humans) as well as their writers, erasers and proposed functions during mitosis.
| PTM site | Writer | Eraser | Cell cycle-dependent regulation & function |
|---|---|---|---|
| Histone H2A | |||
| H2AS1ph | unknown | Increased in S and M phase; required for histone deposition in S phase & chromatin condensation in M phase? [36] | |
| H2AT120ph | Bub1 | Increased at mitotic centromeres; required for recruitment of Sgo1 and Aurora B [35,63] | |
| Histone H2B | |||
| H2BS6ph | CDK1-cyclin B1 | PP1 | Increased at inner centromeres during mitosis; SET displacement; important for chromosome segregation [37] |
| Histone H3 | |||
| H3T3ph | Haspin | PP1γ-Repo-Man | Increased during mitosis; required for Aurora B recruitment; allows displacement of PHD finger proteins [27,30,104,168] |
| H3K4me2/3 | SET1, MLL1 | LSD1, JARID1A | Maintained during mitosis; required for centromeric transcription & bookmarking of active genes [5,57] |
| H3T6ph | unknown | Increased during mitosis; allows displacement of PHD finger proteins [30] | |
| H3K9me2/3 | SUV39H1/2 | JMJD2A | Increased at pericentromeres; required for heterochromatin formation & HP1 recruitment [107–109] |
| H3S10ph | Aurora B | PP1γ | Increased during mitosis; HP1 displacement [22,112,113] |
| H3T11ph | unknown | PP1γ | Increased during mitosis at pericentromeric heterochromatin [31] |
| H3K27me3 | EZH2 | UTX, JMJD3 | Maintained during mitosis; heterochromatin bookmarking [5] |
| H3S28ph | Aurora B | PP1γ | Increased during mitosis; displacement of Polycomb group proteins? [25,130] |
| H3K36me2/3 | NSD2, SETD2 | JMJD2A | Maintained during mitosis; required for centromeric transcription & bookmarking of active genes [5,57] |
| H3K79me2 | DOT1L | KDM2B | Increased during mitosis; important for chromosomal stability [45] |
| H3T80ph | unknown | Increased during mitosis; important for chromosomal stability [28] | |
| H3T118ph | Aurora A | Increased during mitosis; decreased interaction between DNA and histone core; regulation of condensin I and cohesin binding to chromatin [29] | |
| Histone H4 | |||
| H4S1ph | unknown | Increased in S and M phase; required for histone deposition in S phase & chromatin condensation in M phase? [36] | |
| H4K20me1 | PR-Set7 | PHF8 | Increased during mitosis; required for condensin II binding & CENP-A deposition [46,48,84] |
| Histone variants | |||
| H2A.XS121ph | Aurora B | Increased at mitotic centromeres; Aurora B auto-activation at inner centromeres [40] | |
| H2A.XS139ph (γH2A.X) | ATM, CHK2, DNA-PK | PP2A, PP4 | Detected during mitosis; marks unresolved DNA damage? [42–44] |
| H3.3S31ph | Aurora B | Increased during mitosis; p53 activation in response to chromosome missegregation [39,71,72] | |
| CENP-AG1me3 | NRMT1 | Recruitment of CENP-I and CENP-T [92] | |
| CENP-AS7ph | Aurora A, Aurora B | PP1γ? | Increased during mitosis; Aurora B activation; cohesin protection [41,65,66] |
| CENP-AS16ph | unknown | Important for proper chromosome segregation (together with S18) [68] | |
| CENP-AS18ph | CDK2-cyclin E | Regulation of CENP-A deposition [68,93] | |
| CENP-AS68ph | CDK1-cyclin B1 | PP1α | Inhibition of HJURP-mediated CENP-A deposition during mitosis [88] |
| CENP-AK124ub | CUL4A-RBX1-COPS8 | Mediates interaction with HJURP and deposition at centromeres [96] | |
| Linker histones | |||
| H1S/Tph (multiple sites) | CDKs | Increasing phosphorylation during cell cycle progression; involved in transcription, replication, chromatin compaction [38,161] | |
| H1.4T17ph/H1.5S17ph | CDKs | Increased in mitosis; increased H1 mobility & eviction from chromatin by SET [74,161] | |
| H1.4S27ph | Aurora B | Increased in mitosis; precludes HP1 binding; decreased H1 mobility [114,115] | |
| H1.4S35ph | PKA | Increased in mitosis; mediates H1.4 dissociation from chromatin [163] | |
| H1.5T10ph | GSK-3 | Increased in mitosis; increased H1.5 binding to chromatin [164] |