Table 1.

Transformation-responsive telomeric proteins identified by QTIP, reported functions, and association with cancer.

Gene name	Major pathway	Canonical function	Cancer association
APOA1	Lipid metabolism	Participates in cholesterol transport from tissues to liver for excretion; roles in human sperm motility.	(Wang et al, 2009; Su et al, 2010; van Duijnhoven et al, 2011; Jiang et al, 2014; Zamanian-Daryoush & DiDonato, 2015)
ARHGAP1	Rho GTPase signaling	GTPase activator for the Rho, Rac, and Cdc42 proteins.	(Ahn et al, 2012; Li et al, 2017)
CAT	Antioxidant activity	Decomposition of hydrogen peroxide.	(Bauer, 2012; Glorieux et al, 2015; Wang et al, 2016)
CEP170	Microtubule dynamics	Role in microtubule organization and cell morphology.	Unknown
CLPX	Proteostasis	Component of the mitochondrial unfoldase–peptidase complex.	Unknown (Seo et al, 2016)
CLSTN1	Intracellular transport	Regulates kinesin-mediated cargo transport and organizes microtubule polarity during axon development.	Unknown
CTNND1	Cell adhesion, transcription	Transcription regulation, cell adhesion, Wnt signaling, and spatiotemporal control of small Rho-GTPases.	(Thoreson & Reynolds, 2002; Dann et al, 2014; Kourtidis et al, 2015; Li et al, 2015)
DCLRE1B (Apollo)	DNA repair, telomere maintenance	5′-3′ exonuclease, control of DNA damage repair and topological stress, generation and maintenance of telomeric overhangs, and DNA replication.	Unknown (Natrajan et al, 2007; Karami et al, 2016)
GOPC	Proteostasis	Intracellular protein trafficking and degradation.	(Charest et al, 2003; Ohara et al, 2017)
HDGF	Transcription regulation	Heparin-binding protein with mitogenic activity. Regulates transcription.	(Hu et al, 2003; Chen et al, 2015; Lian et al, 2015; Wu et al, 2016; Yang et al, 2016)
HSD17B11	Lipid metabolism	Converts 5α-androstan-3α, 17β-diol to androsterone.	(Nakamura et al, 2009)
HSPB6	Proteostasis	Acts as a molecular chaperone.	(Noda et al, 2007; Matsushima-Nishiwaki et al, 2011, 2016; Nagasawa et al, 2014; Qiao et al, 2014; Ju et al, 2015)
KIF23	Cytokinesis	Component of the centralspindlin complex, essential for cytokinesis in Rho-mediated signaling.	(Takahashi et al, 2012; Sun et al, 2015, 2016; Kato et al, 2016)
MAP4K4	Protein kinase	Activates several mitogen-activated protein kinase pathways; involved in cancer cell growth, apoptosis, and migration.	(Qiu et al, 2012; Haas et al, 2013; Feng et al, 2016; Liu et al, 2016; Gao et al, 2017)
MCM4, 6, 7	DNA replication	Components of the MCM2–7 DNA replicative helicase.	(Honeycutt et al, 2006; Shima et al, 2007; Bagley et al, 2012; Das et al, 2013; Kwok et al, 2015)
MPRIP	Cytoskeleton regulation	Targets myosin phosphatase to the actin cytoskeleton. Regulation of the actin cytoskeleton by RhoA and ROCK1.	(Ono et al, 2008)
NAV1	Cytoskeleton regulation	Regulation of the microtubule cytoskeleton, important for neuronal development, and interacts with the RhoGEF TRIO.	Unknown
NPAT	Transcription, cell cycle	Required for progression through the G1 and S phases, for S phase entry, and for activation of histone gene transcription.	(Saarinen et al, 2011; Hamdi et al, 2017)
ORC1, 3, 5	DNA replication	Components of the origin recognition complex; required for the assembly of pre-RC.	Unknown (Champeris Tsaniras et al, 2014)
PDLIM5	Calcium signaling	Regulates intracellular calcium levels by linking calcium channel and PKC.	(Eeles et al, 2009; Kim et al, 2010)
PKP4	Cell adhesion	Regulates RhoA signaling during cytokinesis.	Unknown
POLDIP2	DNA replication and repair	DNA replication and damage repair, also implicated in mitochondrial function, extracellular matrix regulation, cell cycle progression, focal adhesion, and cell migration.	(Grinchuk et al, 2010; Chian et al, 2016)
RAI14	Cytoskeleton regulation	May have roles in human testis development and spermatogenesis.	Unknown
RNMTL1	RNA processing	Catalyses the formation of 2′-o-methylguanosine at position 1370 in the 16S mitochondrial large-subunit rRNA.	(Haiman et al, 2013)
SAMHD1	dNTP and nucleic acid metabolism	dNTPase activity, has roles in DNA repair, innate immunity, cancer, and HIV-1 restriction; controversial nuclease activity.	(Clifford et al, 2014; Shi et al, 2014; Wang et al, 2014; de Silva et al, 2014; Merati et al, 2015; Rentoft et al, 2016)
SPTAN1	Regulation of cytoskeleton	A scaffolding protein involved in cell adhesion and motility; implicated in repair of DNA interstrand cross-links.	(Tuominen et al, 1996; Gorman et al, 2007; Cunha et al, 2010; Wolgast et al, 2011; Hinrichsen et al, 2014)
SPTBN1	Regulation of cytoskeleton	Scaffolding function in protein sorting, cell adhesion, and migration; implicated in TGF-β signaling.	(Gorman et al, 2007; Jiang et al, 2010; Yao et al, 2010; Zhi et al, 2015)
TCEA1	Transcription	Required for efficient RNA polymerase II transcription elongation past template-encoded arresting sites.	(Hubbard et al, 2008; Shema et al, 2011)
TERF2	Telomere maintenance	Required for telomere capping and protection. Inhibits nonhomologous end joining and ATM activation at telomeres. Required for t loop formation.	(Nakanishi et al, 2003; Bellon et al, 2006; Munoz et al, 2006; Diehl et al, 2011; Biroccio et al, 2013; Pal et al, 2015; Chen et al, 2017)
TMPO	Nuclear architecture	Involved in the structural organization of the nucleus and in the post-mitotic nuclear assembly.	(Brachner & Foisner, 2014; Zhang et al, 2016)
TP53	DDR, cell cycle	Transcription factor inducing cell cycle arrest/senescence and apoptosis; induction of DNA repair genes.	(Muller et al, 2011; Muller & Vousden, 2013; Kaiser & Attardi, 2017)
TPR	Nucleocytoplasmic transport	Scaffolding element of the nuclear pore complex essential for normal nucleocytoplasmic transport of proteins and mRNAs.	(David-Watine, 2011)
TRIOBP	Cytoskeleton regulation	Might be involved in actin remodeling, directed cell movement, and cell cycle regulation.	(Bao et al, 2015; Thutkawkorapin et al, 2016)
TRMT6, TRMT61A	tRNA modification	Catalyses the formation of N1-methyladenine at position 58 in initiator methionyl-tRNA.	(Shi et al, 2015; Macari et al, 2016)