TABLE 3.

Members of the CCP family that act as deglutamylases.

Enzyme and Homologs	PTM	Substrate Preference	Subcellular/Ciliary Distribution	References
CCP1/Nan1 CCPP-1 (Caenorhabditis elegans)	Deglutamylation Δ2 modification Δ3 modification	Detyrosinated α-tubulin PolyE side chain Branching point E Non-Tubulin Protein(s): MLCK-1, Telokin, Klf4, 40S RPS9, TRAD1, HMGB1/2/3	Cilia Cell body (excluded from the nucleus) Dendrites §Also found in the nucleus of HeLa cells	Kalinina et al., 2007; Rogowski et al., 2010; O’Hagan et al., 2011; Berezniuk et al., 2012; De La Vega Otazo et al., 2013; Tort et al., 2014; Tanco et al., 2015; Bompard et al., 2018; Ye et al., 2018
CCP2	Deglutamylation Δ2 modification §No detyrosination or deglycylation	Detyrosinated α-tubulin Poly E side chain	Centrioles Basal bodies Cell body (excluded from the nucleus)	Kalinina et al., 2007; De La Vega Otazo et al., 2013; Tort et al., 2014
CCP3	Deglutamylation Δ2 modification Deaspartylation §No detyrosination or deglycylation	Detyrosinated α-tubulin Poly E side chain	Cell body (excluded from the nucleus)	Kalinina et al., 2007; Tort et al., 2014
CCP4	Deglutamylation Δ2 modification	Detyrosinated α-tubulin PolyE side chain Non-Tubulin Protein(s): MLCK-1, Telokin	Cell body (excluded from the nucleus)	Kalinina et al., 2007; Rogowski et al., 2010
CCP5/Agbl5 *CCPP-6 (Caenorhabditis elegans)	Deglutamylation	Branching point E Poly E side chain (short)	Cilia Cell body Nucleus Mitotic spindle microtubules Midbodies §Cell cycle–dependent distribution: in the nucleus during interphase; in mitotic spindle microtubules, midbodies during mitosis	Kalinina et al., 2007; Kimura et al., 2010; Rogowski et al., 2010; Ghosh-Roy et al., 2012; Berezniuk et al., 2013; De La Vega Otazo et al., 2013; Wu et al., 2017; He et al., 2018
CCP6	Deglutamylation Δ2 modification	Detyrosinated α-tubulin Poly E side chain (long) Non-Tubulin Protein(s): MLCK-1, Telokin, Klf4, Mad2	Cell body (excluded from the nucleus) Basal bodies Golgi apparatus Centrioles §Cell cycle–dependent distribution: in the Golgi apparatus, centrioles during interphase; in centrioles during mitosis	Kalinina et al., 2007; Rogowski et al., 2010; De La Vega Otazo et al., 2013; Ye et al., 2014, 2018

*There is no CCP5 in C. elegans, which do not possess motile cilia. CCPP-6 is functionally similar to CCP5 and is therefore taken to be the ortholog of CCP5.