Table 1.
Mitotic roles of transcription factors.
Name (Organism) |
Mitotic Localization |
Interacting Mitotic Proteins | Loss-of-Function Mitotic Phenotype | Mitotic Function (Role in Cancer) |
---|---|---|---|---|
Sak1, Fkh2 (Sp) | ND | ND | Regulate mitotic gene transcription. Defective septation [24]. | Indirect |
FKHRL1/FOXO3 (h) | ND | ND | Regulates mitotic gene transcription. Defective mitotic exit and cytokinesis [23]. | Indirect (TS) |
Zas1 (Sp) | Chromosomes | ND | Regulates mitotic gene transcription. Defective chromosome condensation and segregation [25]. | Indirect |
ERG (h) | RNA processing bodies (PBs) |
ND | Degrades AURKA and AURKB mRNAs. Multipolar spindles and aberrant spindle structure [26]. | Indirect (TP) |
BRCA1 (h) | Not relevant | Not relevant | Regulates transcription of multiple mitotic genes, including BUB1, BUBR1/BUB1B, AURKA, ESPL1, PTTG1, ASPM, PRC1, and PLK1 [99]. | Indirect (TS) |
BRCA1 (m) | Not relevant | Not relevant | Regulates SAC gene transcription, including MAD2, BUB1, BUBR1, and ZW10 [54,100]. | Indirect (TS) |
BRCA1 (h) | Centrosomes | γ-tubulin | Centrosome amplification and fragmentation [31,32,33,34,35,36,37]. | DIRECT (TS) |
BRCA1 (Xe) | Spindle poles | TPX2, NuMa, XRHAMM | Morphologically abnormal spindles in BRCA1-immunodepleted Xenopus egg extracts [40]. | DIRECT |
ATF5 (h) | Centrosomes (mother centriole) Midbody DZ |
γ-tubulin, PCNT |
Defective accumulation of PCM; centriolar fragmentation, multipolar spindles; unknown function at the midbody [44,45]. | DIRECT |
AKNA (m) | Centrosomes (mother centriole) |
γTuRC, EB1, DCTN1 | Defective centrosome-driven MT regrowth after MT depolymerization [49]. | DIRECT |
YB-1/YBX1 (h) | Centrosomes Spindle poles |
γ-tubulin, PCNT | Structurally abnormal centrosomes with reduced MT nucleation ability [50,51]. | DIRECT (TP) |
OCT1/POU2F1 (h) Interacts with BRCA1 |
Centrosomes Kinetochores Midbody DZ |
PARP-1, APC1 |
Abnormal mitosis; unknown function at the midbody [52,53,54,55]. | DIRECT (TP) |
Oct1 (Xe) | ND | ND | Morphologically abnormal spindles in Oct1-immunodepleted Xenopus egg extracts [52]. | DIRECT |
Nubbin (Dm) (OCT1 ortholog) |
Central spindle Midbody FR |
ND | Unknown function at the midbody [56]. | P-direct ND |
p53/TP53 (h) | Centrosomes | ND | Centrosome amplification and fragmentation [60,61,65,66,67]. | P-direct (TS) |
p53 (m) | Centrosomes | ND | Centrosome amplification [58,62,63,64]. | P-direct |
SP1 (h) | Centrosomes | ND | Centrosome amplification and decreased centrosome-driven MT nucleation [68]. | P-direct (TP) |
SP1 (m) | Centrosomes | ND | Centrosome amplification and decreased centrosome-driven MT nucleation [68]. | P-direct |
SF-1/NR5A1 (m) | Centrosomes | ND | Centrosome amplification [69,70]. | P-direct |
Kaiso/ZBTB33 (h) | Centrosomes Spindle Central spindle Midbody FR |
γ-tubulin, PCNT |
No obvious mitotic defects [71,72]. | P-direct MR (TS) |
CTCF (h) Interacts with Kaiso |
Centrosomes Spindle poles Midbody DZ |
ND | Mitotic phenotype not investigated [73,74]. | P-direct ND (TS) |
SNAP45/SNAPC2 (h) | Centrosomes Spindle poles Midbody FR |
ND | Multiple mitotic defects; defective chromosome condensation [75]. | P-direct |
ELK1 (h) | Centrosomes Spindle Central spindle Midbody FR |
AURKA | No obvious mitotic defects [20,76]. | P-direct MR (TP, TS) |
RUNX1 (h) | Not relevant | Not relevant | Regulates transcription of BUBR1, BUB1, and NEK6 [78]. | Indirect (TP) |
RUNX1, RUNX2, RUNX3 (h) | Centrosomes Spindle Midbody FR |
γ-tubulin, rootletin |
Reduced cyclin B1 accumulation and delayed mitotic entry. No specific mitotic defects [79,80]. | P-direct MR (TP) |
CBFβ/CBFB (h) Binds RUNX proteins |
Midbody FR | MRLC3 | Defective midbody structure and cytokinesis (abscission); polyploid cells [88]. | DIRECT (TS, TP) |
YAP/YAP1 (h) Binds RUNX proteins |
Midbody DZ | PATJ | Defective cytokinesis (abscission) [90]. | DIRECT (TS, TP) |
Aire (m) (AIRE in humans) |
Spindle Central spindle |
AURKB, CEP55, CNTROB, HAUS5, HAUS8, CLASP1, CLASP2 | Abnormal spindle poles and centrosome amplification [94]. | DIRECT |
HDAC3 (h) (in complex with NCOR1, TBL1X and TBL1XR1) |
Spindle | ND | Morphologically abnormal spindles and chromosome misalignment [96]. | P-direct (TP) |
Egr3 (m) | Meiotic spindles of mouse females | ND | Meiotic phenotype not investigated [97]. | P-direct ND |
TFIIB (m) (GTF2B in humans) |
Meiotic spindles of mouse females | ND | Morphologically abnormal spindles and chromosomes misalignment after antibody injection [98]. | P-direct |
WT1 (h) | Kinetochores (colocalizes with MAD2) |
MAD2 | Accelerated metaphase-to-anaphase transition; defective chromosome segregation [101,102]. | DIRECT (TS, TP) |
TIF1γ/TRIM33 (h) | ND |
CDC20, APC/C |
Chromosome misalignment; delay in metaphase-to-anaphase transition [103,104]. | P-direct (TS, TP) |
XPB/ERCC3 (h) (TFIIH subunit) |
Centrosomes Spindle poles |
γ-tubulin | Mitotic phenotype not investigated [111]. | Direct ND (TS) |
Hay (Dm) (TFIIH subunit, orthologous to XPB) |
Chromosome/ spindle area in embryos |
Testis-specific β2-tubulin | Defective meiotic spindles in males; abnormal mitotic spindles and defective chromosome segregation in embryos from hay mutant mothers [108,109,110,113]. | P-direct |
p8, p52, Cdk7 (Dm) (TFIIH subunits) |
Chromosome/ spindle area in embryos |
ND | Abnormal mitotic spindles and defective chromosome segregation in embryos from mutant or RNAi mothers [113]. | P-direct |
XPD/ERCC2 (h) (TFIIH subunit forms another complex with MIP18 and MMS19) |
Spindle (with MIP18 and MMS19) |
ND | Multipolar spindles and multinucleated cells [112]. | P-direct |
Xpd (Dm) (TFIIH subunit) |
ND | Regulates Cdk7 localization | Abnormal mitotic spindles and defective chromosome segregation in embryos from Xpd mutant mothers [114]. | P-direct |
KANSL1, KANSL3, MCRS1 (h) (KANSL complex subunits) |
Spindle poles; KANSL1 and KANSL3 bind MT minus ends | TPX2, MCAK |
Chromosome misalignment and defective segregation [120,121]. | DIRECT (TP) |
WDR5 (h) (KANSL subunit that forms another complex with MLL1/KMT2A) |
Centrosomes Spindle Central spindle Midbody DZ |
KIF2A, PRC1, MKLP1, CYK4, CEP55 |
Morphologically abnormal and elongated spindles, aberrant chromosome segregation, and failures in cytokinesis [124,125,126]. | DIRECT (TP) |
Dgt1 (Dm) (NSL subunit orthologous to KANSL2) |
Diffuse (weak on spindle) |
ND | Diminished γ-tubulin at centrosomes; long spindles [17]. | P-direct |
Rcd1 (Dm) (NSL subunit orthologous to KANSL3) |
Centrosomes Central spindle Midbody FR |
ND | Defective centriole replication; frequent failures in chromosome alignment and segregation [18,19,127]. | Indirect (P-direct MR) |
Rcd5 (Dm) (NSL subunit orthologous to MCRS1) |
Centrosomes Midbody DZ |
ND | Defective centriole replication; frequent failures in chromosome alignment and segregation [17,18,127]. | Indirect (P-direct MR) |
MBD-R2 (Dm) (NSL subunit orthologous to PHF20) |
Chromosomes | ND | Defective centriole replication; frequent failures in chromosome alignment and segregation [19,127]. | Indirect (P-direct MR) |
Wds (Dm) (NSL subunit orthologous to WDR5) |
Centrosomes Kinetochores Midbody DZ |
ND | Defective centriole replication; frequent failures in chromosome alignment and segregation [127]. | Indirect (P-direct MR) |
Name: Each human TF is designated either with its HUGO acronym only, or both the name used in the cited papers and the HUGO acronym. Organism: h, human; m, mouse; Dm, Drosophila melanogaster, Sp., Schizosaccharomyces pombe, Xe, Xenopus egg extracts. Mitotic localization: ND, not determined; Midbody DZ, midbody dark zone; Midbody FR, midbody flanking regions; localization to specific midbody regions has been decided based on the examination of published microphotographs. Interacting mitotic proteins: ND, not determined. Mitotic function: Indirect, controls transcription of mitotic genes; DIRECT, in addition to a role in transcription has a direct role in mitosis, namely it is a moonlighting protein that meets the 3 criteria specified in the text. P-direct, putatively direct, does not meet the 3 criteria but (in the cited papers) is thought to have a direct mitotic role. P-direct ND, localizes to a mitotic structure but its mitotic role has not been investigated. P-direct MR, localizes to a mitotic structure but its loss does not result in a detectable mitotic phenotype; it might therefore have either a minor or a redundant mitotic function. Role in cancer was determined by examination of the pertinent literature; references are not reported: TS, tumor suppressor; TP, tumor promoter; TS, TP, can function either as a tumor suppressor or promoter, depending on the cellular context.