Q04859 (MAK_MOUSE) Reviewed, UniProtKB/Swiss-Prot

Names and origin

Protein names

Recommended name:
Serine/threonine-protein kinase MAK
EC=2.7.11.22

Alternative name(s):
Male germ cell-associated kinase
Protein kinase RCK

Gene names

Name:	Mak
Synonyms:	Rck

Organism

Mus musculus (Mouse) [Reference proteome]

Taxonomic identifier

10090 [NCBI]

Taxonomic lineage

cellular organisms › Eukaryota › Opisthokonta › Metazoa › Eumetazoa › Bilateria › Deuterostomia › Chordata › Craniata › Vertebrata › Gnathostomata › Teleostomi › Euteleostomi › Sarcopterygii › Tetrapoda › Amniota › Mammalia › Theria › Eutheria › Euarchontoglires › Glires › Rodentia › Sciurognathi › Muroidea › Muridae › Murinae › Mus › Mus

Protein attributes

Sequence length	622 AA.
Sequence status	Complete.
Protein existence	Evidence at protein level

General annotation (Comments)

Function	Essential for the regulation of ciliary length and required for the long-term survival of photoreceptors. Could have an important function in sensory cells and in spermatogenesis. May participate in signaling pathways used in visual and olfactory sensory transduction. Phosphorylates FZR1 in a cell cycle-dependent manner. Plays a role in the transcriptional coactivation of AR By similarity. Ref.6
Catalytic activity	ATP + a protein = ADP + a phosphoprotein.
Subunit structure	Interacts with AR and CDK20. Found in a complex containing MAK, AR and NCOA3. Interacts with FZR1 (via WD repeats) By similarity. Interacts with RP1. Ref.6
Subcellular location	Nucleus By similarity. Cytoplasm › cytoskeleton › centrosome By similarity. Cytoplasm › cytoskeleton › spindle By similarity. Midbody By similarity. Cell projection › cilium › photoreceptor outer segment. Photoreceptor inner segment By similarity. Note: Localizes in both the connecting cilia and the outer segment axonemes. Ref.6
Tissue specificity	In pre- and postmeiotic male germ cells in testis. In photoreceptor cells of the retina and in the olfactory receptors, and in certain epithelia of the respiratory tract and choroid plexus (brain). Ref.6
Developmental stage	On day 14 or 17 of embryonic development. Expression is observed in germ cells at the stages of late pachytene spermatocytes through to early round spermatids. Ref.4
Post-translational modification	Autophosphorylated. Phosphorylated on serine and threonine residues By similarity.
Disruption phenotype	Mice are mostly fertile, develop normally, and exhibit no gross abnormalities and spermatogenesis is intact. However, both sperm motility and litter size is reduced. Ref.5
Sequence similarities	Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. CDC2/CDKX subfamily. Contains 1 protein kinase domain.

Ontologies

Keywords
Biological process	Transcription Transcription regulation
Cellular component	Cell projection Cilium Cytoplasm Cytoskeleton Nucleus
Coding sequence diversity	Alternative splicing
Ligand	ATP-binding Nucleotide-binding
Molecular function	Kinase Serine/threonine-protein kinase Transferase
PTM	Phosphoprotein
Technical term	Complete proteome Reference proteome
Gene Ontology (GO)
Biological_process	cell cycle Inferred from electronic annotation. Source: GOC photoreceptor cell maintenance Inferred from mutant phenotype Ref.6. Source: UniProtKB protein autophosphorylation Inferred from sequence or structural similarity. Source: UniProtKB regulation of transcription, DNA-dependent Inferred from electronic annotation. Source: UniProtKB-KW transcription, DNA-dependent Inferred from electronic annotation. Source: UniProtKB-KW
Cellular_component	centrosome Inferred from sequence or structural similarity. Source: UniProtKB cytoplasm Inferred from electronic annotation. Source: UniProtKB-KW midbody Inferred from sequence or structural similarity. Source: UniProtKB mitotic spindle Inferred from sequence or structural similarity. Source: UniProtKB nucleus Inferred from sequence or structural similarity. Source: UniProtKB photoreceptor inner segment Inferred from sequence or structural similarity. Source: UniProtKB photoreceptor outer segment Inferred from direct assay Ref.6. Source: UniProtKB
Molecular_function	ATP binding Inferred from electronic annotation. Source: UniProtKB-KW cyclin-dependent protein serine/threonine kinase activity Inferred from electronic annotation. Source: EC transcription coactivator activity Inferred from sequence or structural similarity. Source: UniProtKB
Complete GO annotation...

Alternative products

This entry describes 3 isoforms produced by alternative splicing. [Align] [Select]

Isoform 1 (identifier: Q04859-1)

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

Isoform 2 (identifier: Q04859-2)

The sequence of this isoform differs from the canonical sequence as follows:
     531-531: A → EDSIIKPIENLSCTGKSAEQLEDPQ

Isoform 3 (identifier: Q04859-3)

The sequence of this isoform differs from the canonical sequence as follows:
     94-124: Missing.
     531-571: Missing.

Note: No experimental confirmation available.

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Chain	1 – 622	622	Serine/threonine-protein kinase MAK		PRO_0000086285
Regions
	Domain	4 – 284	281	Protein kinase
	Nucleotide binding	10 – 18	9	ATP By similarity
	Compositional bias	309 – 368	60	Glu/Pro-rich
Sites
	Active site	125	1	Proton acceptor By similarity
	Binding site	33	1	ATP By similarity
Amino acid modifications
	Modified residue	157	1	Phosphothreonine; by autocatalysis By similarity
	Modified residue	159	1	Phosphotyrosine; by autocatalysis By similarity
Natural variations
	Alternative sequence	94 – 124	31	Missing in isoform 3.		VSP_042472
	Alternative sequence	531 – 571	41	Missing in isoform 3.		VSP_042473
	Alternative sequence	531	1	A → EDSIIKPIENLSCTGKSAEQ LEDPQ in isoform 2.		VSP_042474
Experimental info
	Sequence conflict	128	1	P → H in BAC26662. Ref.2
	Sequence conflict	235	1	N → D in BAC26662. Ref.2
	Sequence conflict	386	1	T → A in CAA47392. Ref.1

Sequences

	Sequence		Length	Mass (Da)
	Isoform 1 [UniParc]. Last modified July 27, 2011. Version 2. Checksum: DB7E3B758F8FEA69	FASTA	622	70,080
	10 20 30 40 50 60 MNRYTTMKQL GDGTYGSVLM GKSNESGELV AIKRMKRKFY SWDECMNLRE VKSLKKLNHA 70 80 90 100 110 120 NVIKLKEVIR ENDHLYFVFE YMKENLYQLM KDRNKLFPES VIRNIMYQIL QGLAFIHKHG 130 140 150 160 170 180 FFHRDMKPEN LLCMGPELVK IADFGLAREL RSQPPYTDYV STRWYRAPEV LLRSSVYSSP 190 200 210 220 230 240 IDVWAVGSIM AELYTFRPLF PGTSEVDEIF KICQVLGTPK KSDWPEGYQL ASSMNFRFPQ 250 260 270 280 290 300 CIPINLKTLI PNASSEAIQL MTEMLNWDPK KRPTASQALK HPYFQVGQVL GSSAHHLDTK 310 320 330 340 350 360 QTLHKQLQPL EPKPSSSERD PKPLPNILDQ PAGQPQPKQG HQPLQTIQPP QNTVTHPPPK 370 380 390 400 410 420 QQGHQKPPQT MFPSIIKTIP VNSVSTLGHK GARRRWGQTV FKSGDSCDDI EDDLGASHSK 430 440 450 460 470 480 KPSMEACKEK KKESPFRFPD SGLPVSNHFK GENRNLHASV SLKSDPNLST ASTAKQYYLK 490 500 510 520 530 540 QSRYLPGVNP KNVSLVAGGK DINSHSWNNQ LFPKSLGSMG ADLSFKRSNA AGNLGSYTTY 550 560 570 580 590 600 NQTGYMPSFL KKEVGSAGQR IQLAPLGASA SDYTWSTKTG RGQFSGRTYN PTAKNLNIVN 610 620 RTQPVPSVHG RTDWVAKYGG HR
	Isoform 2 [UniParc]. Checksum: 05CA06912663F2CC	FASTA	646	72,736
	10 20 30 40 50 60 MNRYTTMKQL GDGTYGSVLM GKSNESGELV AIKRMKRKFY SWDECMNLRE VKSLKKLNHA 70 80 90 100 110 120 NVIKLKEVIR ENDHLYFVFE YMKENLYQLM KDRNKLFPES VIRNIMYQIL QGLAFIHKHG 130 140 150 160 170 180 FFHRDMKPEN LLCMGPELVK IADFGLAREL RSQPPYTDYV STRWYRAPEV LLRSSVYSSP 190 200 210 220 230 240 IDVWAVGSIM AELYTFRPLF PGTSEVDEIF KICQVLGTPK KSDWPEGYQL ASSMNFRFPQ 250 260 270 280 290 300 CIPINLKTLI PNASSEAIQL MTEMLNWDPK KRPTASQALK HPYFQVGQVL GSSAHHLDTK 310 320 330 340 350 360 QTLHKQLQPL EPKPSSSERD PKPLPNILDQ PAGQPQPKQG HQPLQTIQPP QNTVTHPPPK 370 380 390 400 410 420 QQGHQKPPQT MFPSIIKTIP VNSVSTLGHK GARRRWGQTV FKSGDSCDDI EDDLGASHSK 430 440 450 460 470 480 KPSMEACKEK KKESPFRFPD SGLPVSNHFK GENRNLHASV SLKSDPNLST ASTAKQYYLK 490 500 510 520 530 540 QSRYLPGVNP KNVSLVAGGK DINSHSWNNQ LFPKSLGSMG ADLSFKRSNA EDSIIKPIEN 550 560 570 580 590 600 LSCTGKSAEQ LEDPQGNLGS YTTYNQTGYM PSFLKKEVGS AGQRIQLAPL GASASDYTWS 610 620 630 640 TKTGRGQFSG RTYNPTAKNL NIVNRTQPVP SVHGRTDWVA KYGGHR
	Isoform 3 [UniParc]. Checksum: 639D4B5A3A7465B0	FASTA	550	62,125
	10 20 30 40 50 60 MNRYTTMKQL GDGTYGSVLM GKSNESGELV AIKRMKRKFY SWDECMNLRE VKSLKKLNHA 70 80 90 100 110 120 NVIKLKEVIR ENDHLYFVFE YMKENLYQLM KDRDMKPENL LCMGPELVKI ADFGLARELR 130 140 150 160 170 180 SQPPYTDYVS TRWYRAPEVL LRSSVYSSPI DVWAVGSIMA ELYTFRPLFP GTSEVDEIFK 190 200 210 220 230 240 ICQVLGTPKK SDWPEGYQLA SSMNFRFPQC IPINLKTLIP NASSEAIQLM TEMLNWDPKK 250 260 270 280 290 300 RPTASQALKH PYFQVGQVLG SSAHHLDTKQ TLHKQLQPLE PKPSSSERDP KPLPNILDQP 310 320 330 340 350 360 AGQPQPKQGH QPLQTIQPPQ NTVTHPPPKQ QGHQKPPQTM FPSIIKTIPV NSVSTLGHKG 370 380 390 400 410 420 ARRRWGQTVF KSGDSCDDIE DDLGASHSKK PSMEACKEKK KESPFRFPDS GLPVSNHFKG 430 440 450 460 470 480 ENRNLHASVS LKSDPNLSTA STAKQYYLKQ SRYLPGVNPK NVSLVAGGKD INSHSWNNQL 490 500 510 520 530 540 FPKSLGSMGA DLSFKRSNAD YTWSTKTGRG QFSGRTYNPT AKNLNIVNRT QPVPSVHGRT 550 DWVAKYGGHR

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"Characterization and expression analysis of the murine rck gene: a protein kinase with a potential function in sensory cells." Bladt F., Birchmeier C. Differentiation 53:115-122(1993) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 1).
[2]	"The transcriptional landscape of the mammalian genome." Carninci P., Kasukawa T., Katayama S., Gough J., Frith M.C., Maeda N., Oyama R., Ravasi T., Lenhard B., Wells C., Kodzius R., Shimokawa K., Bajic V.B., Brenner S.E., Batalov S., Forrest A.R., Zavolan M., Davis M.J. , Wilming L.G., Aidinis V., Allen J.E., Ambesi-Impiombato A., Apweiler R., Aturaliya R.N., Bailey T.L., Bansal M., Baxter L., Beisel K.W., Bersano T., Bono H., Chalk A.M., Chiu K.P., Choudhary V., Christoffels A., Clutterbuck D.R., Crowe M.L., Dalla E., Dalrymple B.P., de Bono B., Della Gatta G., di Bernardo D., Down T., Engstrom P., Fagiolini M., Faulkner G., Fletcher C.F., Fukushima T., Furuno M., Futaki S., Gariboldi M., Georgii-Hemming P., Gingeras T.R., Gojobori T., Green R.E., Gustincich S., Harbers M., Hayashi Y., Hensch T.K., Hirokawa N., Hill D., Huminiecki L., Iacono M., Ikeo K., Iwama A., Ishikawa T., Jakt M., Kanapin A., Katoh M., Kawasawa Y., Kelso J., Kitamura H., Kitano H., Kollias G., Krishnan S.P., Kruger A., Kummerfeld S.K., Kurochkin I.V., Lareau L.F., Lazarevic D., Lipovich L., Liu J., Liuni S., McWilliam S., Madan Babu M., Madera M., Marchionni L., Matsuda H., Matsuzawa S., Miki H., Mignone F., Miyake S., Morris K., Mottagui-Tabar S., Mulder N., Nakano N., Nakauchi H., Ng P., Nilsson R., Nishiguchi S., Nishikawa S., Nori F., Ohara O., Okazaki Y., Orlando V., Pang K.C., Pavan W.J., Pavesi G., Pesole G., Petrovsky N., Piazza S., Reed J., Reid J.F., Ring B.Z., Ringwald M., Rost B., Ruan Y., Salzberg S.L., Sandelin A., Schneider C., Schoenbach C., Sekiguchi K., Semple C.A., Seno S., Sessa L., Sheng Y., Shibata Y., Shimada H., Shimada K., Silva D., Sinclair B., Sperling S., Stupka E., Sugiura K., Sultana R., Takenaka Y., Taki K., Tammoja K., Tan S.L., Tang S., Taylor M.S., Tegner J., Teichmann S.A., Ueda H.R., van Nimwegen E., Verardo R., Wei C.L., Yagi K., Yamanishi H., Zabarovsky E., Zhu S., Zimmer A., Hide W., Bult C., Grimmond S.M., Teasdale R.D., Liu E.T., Brusic V., Quackenbush J., Wahlestedt C., Mattick J.S., Hume D.A., Kai C., Sasaki D., Tomaru Y., Fukuda S., Kanamori-Katayama M., Suzuki M., Aoki J., Arakawa T., Iida J., Imamura K., Itoh M., Kato T., Kawaji H., Kawagashira N., Kawashima T., Kojima M., Kondo S., Konno H., Nakano K., Ninomiya N., Nishio T., Okada M., Plessy C., Shibata K., Shiraki T., Suzuki S., Tagami M., Waki K., Watahiki A., Okamura-Oho Y., Suzuki H., Kawai J., Hayashizaki Y. Science 309:1559-1563(2005) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 3). Strain: C57BL/6J. Tissue: Testis.
[3]	"Lineage-specific biology revealed by a finished genome assembly of the mouse." Church D.M., Goodstadt L., Hillier L.W., Zody M.C., Goldstein S., She X., Bult C.J., Agarwala R., Cherry J.L., DiCuccio M., Hlavina W., Kapustin Y., Meric P., Maglott D., Birtle Z., Marques A.C., Graves T., Zhou S. , Teague B., Potamousis K., Churas C., Place M., Herschleb J., Runnheim R., Forrest D., Amos-Landgraf J., Schwartz D.C., Cheng Z., Lindblad-Toh K., Eichler E.E., Ponting C.P. PLoS Biol. 7:E1000112-E1000112(2009) [PubMed] [Europe PMC] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. Strain: C57BL/6J.
[4]	"In situ localization of male germ cell-associated kinase (mak) mRNA in adult mouse testis: specific expression in germ cells at stages around meiotic cell division." Koji T., Jinno A., Matsushime H., Shibuya M., Nakane P.K. Cell Biochem. Funct. 10:273-279(1992) [PubMed] [Europe PMC] [Abstract] Cited for: DEVELOPMENTAL STAGE.
[5]	"A testicular germ cell-associated serine-threonine kinase, MAK, is dispensable for sperm formation." Shinkai Y., Satoh H., Takeda N., Fukuda M., Chiba E., Kato T., Kuramochi T., Araki Y. Mol. Cell. Biol. 22:3276-3280(2002) [PubMed] [Europe PMC] [Abstract] Cited for: DISRUPTION PHENOTYPE.
[6]	"Negative regulation of ciliary length by ciliary male germ cell-associated kinase (Mak) is required for retinal photoreceptor survival." Omori Y., Chaya T., Katoh K., Kajimura N., Sato S., Muraoka K., Ueno S., Koyasu T., Kondo M., Furukawa T. Proc. Natl. Acad. Sci. U.S.A. 107:22671-22676(2010) [PubMed] [Europe PMC] [Abstract] Cited for: FUNCTION, SUBCELLULAR LOCATION, INTERACTION WITH RP1, TISSUE SPECIFICITY.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	X66983 mRNA. Translation: CAA47392.1. AK029894 mRNA. Translation: BAC26662.1. AC133496 Genomic DNA. No translation available.
IPI	IPI00111032. IPI00407263. IPI00928396.
PIR	I48733.
RefSeq	NP_001139274.1. NM_001145802.1. NP_001139275.1. NM_001145803.1. NP_032573.2. NM_008547.2.
UniGene	Mm.8149.
3D structure databases
ProteinModelPortal	Q04859.
SMR	Q04859. Positions 1-353.
ModBase	Search...
Protein-protein interaction databases
DIP	DIP-59494N.
PTM databases
PhosphoSite	Q04859.
Proteomic databases
PaxDb	Q04859.
PRIDE	Q04859.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
Ensembl	ENSMUST00000021792; ENSMUSP00000021792; ENSMUSG00000021363. ENSMUST00000070193; ENSMUSP00000064750; ENSMUSG00000021363. ENSMUST00000165087; ENSMUSP00000129615; ENSMUSG00000021363.
GeneID	17152.
KEGG	mmu:17152.
UCSC	uc007qev.2. mouse.
Organism-specific databases
CTD	4117.
MGI	MGI:96913. Mak.
Phylogenomic databases
eggNOG	COG0515.
GeneTree	ENSGT00650000093283.
HOGENOM	HOG000233024.
HOVERGEN	HBG014652.
InParanoid	Q04859.
KO	K08829.
OMA	NLGSYAT.
Enzyme and pathway databases
BRENDA	2.7.11.22. 3474.
Gene expression databases
Bgee	Q04859.
CleanEx	MM_MAK.
Genevestigator	Q04859.
GermOnline	ENSMUSG00000021363. Mus musculus.
Family and domain databases
InterPro	IPR011009. Kinase-like_dom. IPR000719. Prot_kinase_cat_dom. IPR017441. Protein_kinase_ATP_BS. IPR002290. Ser/Thr_dual-sp_kinase_dom. IPR008271. Ser/Thr_kinase_AS. [Graphical view]
Pfam	PF00069. Pkinase. 1 hit. [Graphical view]
SMART	SM00220. S_TKc. 1 hit. [Graphical view]
SUPFAM	SSF56112. Kinase_like. 1 hit.
PROSITE	PS00107. PROTEIN_KINASE_ATP. 1 hit. PS50011. PROTEIN_KINASE_DOM. 1 hit. PS00108. PROTEIN_KINASE_ST. 1 hit. [Graphical view]
ProtoNet	Search...
Other
NextBio	291410.
SOURCE	Search...

Entry information

Entry name

MAK_MOUSE

Accession

Primary (citable) accession number: Q04859
Secondary accession number(s): E9Q4B2 , E9QAU5, E9QKR3, Q8CDL5

Entry history

Integrated into UniProtKB/Swiss-Prot:	February 1, 1994
Last sequence update:	July 27, 2011
Last modified:	May 1, 2013
This is version 98 of the entry and version 2 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation program

Chordata Protein Annotation Program