Table 1.

Histone residues that are phosphorylateda

HistoneResidueOrganismKinase(s)Protein(s) recognizing (bound or released) the modificationFunction(s)
H2ASerine 139 (H2AX)MammalsATR, ATM, DNA-PK, RSK2, MSK1DSB-sensing proteins (MRN, MDC1, P53BP1), AP1DNA repair, decreased EGF-mediated cellular transformation, apoptosis
Tyrosine 142 (H2AX)MammalsWSTFNDDecision between cell survival and apoptosis
Serine 1MammalsMSKNDInhibition of transcription
Threonine 119MammalsNHK-1, Aurora BPTB domain-containing protein Fe65Regulation of chromatin structure and function during mitosis
Serine 121Fission yeastBub1ShugoshinMaintaining chromosomal homeostasis by recruiting shugoshin at sister kinetochore
Serine 129YeastATM-related kinase Mec1 and Tel1NuA4, SWR1, INO80, cohesinDSB repair
H2BSerine 10YeastSte20NDApoptosis
Serine 14Various vertebratesMstINDApoptosis
Serine 32MammalsProtein kinase CNDPossible involvement in apoptosis-related nucleosomal DNA fragmentation
Serine 33DrosophilaCTK-TAF1NDTranscriptional regulation
Serine 36MammalsAMPKNDDirect transcriptional and chromatin regulatory pathways leading to cellular response to stress
H3Threonine 3MammalsHaspinSurvivinCorrect localization of the CPC at the centromere
Threonine 6MammalsPKCβ1NDAndrogen-dependent H3T6 phosphorylation prevents LSD1-mediated H3K4 demethylation, maintaining hormone-dependent gene activation
Serine 10Yeast, mammalsSnf1, IpL1 (yeast), Aurora B (mammals), MSK1/2, IKKα, PKB/Akt, Rsk2, PIM1HP1, SRp20, ASF/SF2, 14-3-3Help in chromosome condensation during mitosis and meiosis; roles in transcription of certain genes
Threonine 11MammalsChk1, PRK1, Dlk/Zip kinaseGCN5Transcriptional activation of certain genes; probable role at centromeres during mitosis
Serine 28MammalsAurora B, MSK1/2Polycomb silencing complexHelp in chromosome condensation during mitosis and meiosis; roles in transcription of certain genes
Tyrosine 41MammalsJAK2HP1αRole in differentiation related to hematopoiesis
Threonine 45Mammals, budding yeastProtein kinase C, S-phase kinase Cdc7-Dbf4NDApoptosis, role in DNA-damaged cells when the DNA is nicked, replication of DNA
H4Serine 1Yeast, mammalsCK IINDDNA damage repair, mitosis and chromatin assembly
Histidines 18 and 75MammalsUnknownNDFacilitation of DNA replication by destabilizing histone octamer
  • a DSB, double-strand break; ND, not determined.