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protein coding gene - alp4 (SPBC365.15) - gamma tubulin complex GCP2 subunit Alp4

Gene summary

Standard name
alp4
Systematic ID
SPBC365.15
Product
gamma tubulin complex GCP2 subunit Alp4
Organism
Schizosaccharomyces pombe (fission yeast)
UniProt ID
Q9Y705
ORFeome ID
38/38C08
Characterisation status
biological role published
Feature type
mRNA gene
Genomic location
chromosome II: 2524061..2527134 forward strand

Annotation

Complementation

PBO:0091789 - functionally complemented by human GCP2

References:

Disease association

MONDO:0032893 - pachygyria, microcephaly, developmental delay, and dysmorphic facies, with or without seizures

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GO biological process

GO:1990734 - astral microtubule anchoring at mitotic spindle pole body

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GO:0031122 - cytoplasmic microtubule organization

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GO:0051321 - meiotic cell cycle

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GO:0007020 - microtubule nucleation

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GO:0090307 - mitotic spindle assembly

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GO:0098863 - nuclear migration by microtubule mediated pushing forces

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GO cellular component

GO:0032153 - cell division site

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GO:0005829 - cytosol

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GO:0000923 - equatorial microtubule organizing center

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GO:0000931 - gamma-tubulin ring complex

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GO:0008275 - gamma-tubulin small complex

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GO:0061496 - half bridge of mitotic spindle pole body

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GO:0061497 - inner plaque of mitotic spindle pole body

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GO:0031021 - interphase microtubule organizing center

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GO:0035974 - meiotic spindle pole body

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GO:0044732 - mitotic spindle pole body

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GO:0005634 - nucleus

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GO:0071957 - old mitotic spindle pole body

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GO molecular function

GO:0051011 - microtubule minus-end binding

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GO:0140490 - microtubule nucleator activity

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GO:0005515 - protein binding

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Modification

MOD:00046 - O-phospho-L-serine

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MOD:01149 - sumoylated lysine

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Multi-locus phenotype

FYPO:0001355 - decreased vegetative cell population growth

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Genotypes:

FYPO:0004950 - elongated vegetative cell with elongated oscillating nucleus

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Genotypes:

FYPO:0001837 - increased duration of protein localization to mitotic spindle pole body

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Genotypes:

FYPO:0004922 - inviable elongated mononucleate aseptate cell with cell cycle arrest at mitotic G2/M phase transition

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Genotypes:

FYPO:0000276 - monopolar mitotic spindle

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Genotypes:

FYPO:0000899 - normal microtubule cytoskeleton organization during vegetative growth

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Genotypes:

FYPO:0001357 - normal vegetative cell population growth

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Genotypes:

FYPO:0002058 - viable cell population

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Genotypes:

FYPO:0006822 - viable small vegetative cell with normal cell growth rate

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Genotypes:

Qualitative gene expression

PomGeneEx:0000012 - RNA level decreased

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PomGeneEx:0000011 - RNA level increased

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Quantitative gene expression

PBO:0006310 - protein level

References:

PBO:0011963 - RNA level

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Single locus phenotype

FYPO:0000608 - abnormal cell cycle arrest in mitotic M phase

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Genotypes:

FYPO:0003972 - abnormal exit from mitosis

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Genotypes:

FYPO:0004590 - abnormal gamma-tubulin complex localization to mitotic spindle pole body following normal mitosis

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Genotypes:

FYPO:0001350 - abnormal microtubule cytoskeleton organization during vegetative growth

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Genotypes:

FYPO:0000141 - abnormal mitotic sister chromatid segregation

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Genotypes:

FYPO:0000177 - abnormal mitotic spindle assembly

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Genotypes:

FYPO:0000131 - abnormal mitotic spindle elongation

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Genotypes:

FYPO:0004613 - abolished microtubule attachment to spindle pole body

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Genotypes:

FYPO:0004318 - abolished mitotic spindle assembly checkpoint

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Genotypes:

FYPO:0004732 - astral microtubules present in decreased numbers

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Genotypes:

FYPO:0004700 - bent vegetative cell

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Genotypes:

FYPO:0000446 - cell cycle arrest at mitotic G2/M phase transition

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Genotypes:

FYPO:0000016 - curved vegetative cell

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Genotypes:

FYPO:0003165 - cut with abnormal chromosome segregation

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Genotypes:

FYPO:0004733 - cytoplasmic microtubules nucleated from eMTOC present in decreased numbers

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Genotypes:

FYPO:0001875 - decreased asymmetric protein localization, with protein localized to both mitotic spindle pole bodies

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Genotypes:

FYPO:0001876 - decreased asymmetric protein localization, with protein localized to both mitotic spindle pole bodies during anaphase

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Genotypes:

FYPO:0005029 - decreased gamma-tubulin complex localization to mitotic spindle pole body

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Genotypes:

FYPO:0001586 - decreased protein localization to cell tip during vegetative growth

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Genotypes:

FYPO:0004980 - decreased protein localization to eMTOC during vegetative growth

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Genotypes:

FYPO:0000940 - decreased protein localization to mitotic spindle pole body

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Genotypes:

FYPO:0004975 - decreased rate of microtubule depolymerization behind moving spindle pole body

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Genotypes:

FYPO:0004979 - elongated monoseptate vegetative cell

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Genotypes:

FYPO:0004750 - elongated septated vegetative cell

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Genotypes:

FYPO:0001122 - elongated vegetative cell

References:

Genotypes:

FYPO:0004950 - elongated vegetative cell with elongated oscillating nucleus

References:

Genotypes:

FYPO:0005002 - elongated vegetative cell with oscillating nucleus

References:

Genotypes:

FYPO:0000741 - increased cellular pigment accumulation

References:

Genotypes:

FYPO:0001837 - increased duration of protein localization to mitotic spindle pole body

References:

Genotypes:

FYPO:0002852 - increased protein localization to cell tip during vegetative growth

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Genotypes:

FYPO:0002870 - increased protein localization to non-growing cell tip

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Genotypes:

FYPO:0004976 - increased rate of microtubule depolymerization ahead of moving spindle pole body

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Genotypes:

FYPO:0002113 - inviable curved vegetative cell

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Genotypes:

FYPO:0002066 - inviable elongated aseptate cell

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Genotypes:

FYPO:0002337 - inviable elongated mononucleate vegetative cell with mislocalized nucleus

References:

Genotypes:

FYPO:0001489 - inviable vegetative cell

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Genotypes:

FYPO:0002061 - inviable vegetative cell population

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Genotypes:

FYPO:0004511 - long curved interphase microtubules

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Genotypes:

FYPO:0000233 - long cytoplasmic microtubules

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Genotypes:

FYPO:0004537 - mitotic spindle assembly checkpoint override

References:

Genotypes:

FYPO:0000276 - monopolar mitotic spindle

References:

Genotypes:

FYPO:0000118 - multiseptate vegetative cell

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Genotypes:

FYPO:0002002 - multiseptate vegetative cell, septa grouped

References:

Genotypes:

FYPO:0001870 - normal centromere clustering at nuclear periphery during vegetative growth

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Genotypes:

FYPO:0004367 - normal mitotic spindle assembly

References:

Genotypes:

FYPO:0003762 - normal mitotic spindle assembly checkpoint

References:

Genotypes:

FYPO:0001587 - normal protein localization to cell tip during vegetative growth

References:

Genotypes:

FYPO:0004974 - normal telomere localization during mitotic interphase

References:

Genotypes:

FYPO:0002085 - normal vegetative cell growth

References:

Genotypes:

FYPO:0003503 - normal vegetative cell length

References:

Genotypes:

FYPO:0001357 - normal vegetative cell population growth

References:

Genotypes:

FYPO:0003788 - nuclear envelope protrusion present during mitosis

References:

Genotypes:

FYPO:0000091 - sensitive to thiabendazole

References:

Genotypes:

FYPO:0002760 - short cytoplasmic microtubules

References:

Genotypes:

FYPO:0004539 - short interphase microtubules present in increased numbers

References:

Genotypes:

Taxonomic conservation

PBO:0011065 - conserved in eukaryotes

PBO:0011071 - conserved in eukaryotes only

PBO:0011064 - conserved in fungi

PBO:0011069 - conserved in metazoa

PBO:0011070 - conserved in vertebrates

PBO:0006222 - predominantly single copy (one to one)

Protein features

IDNameInterPro nameDB name
PF04130GCP_C_terminalGCP_CPFAM
PF17681GCP_N_terminalGCP_NPFAM
G3DSA:1.20.120.1900:FF:000011FUNFAM
G3DSA:1.20.120.1900GCP_C_sfGENE3D
PTHR19302GAMMA TUBULIN COMPLEX PROTEINGCPPANTHER

Orthologs

References / Literature

PMID:15068790 - Rsp1p, a J domain protein required for disassembly and assembly of microtubule organizing centers during the fission yeast cell cycle.
Zimmerman S et al. Dev Cell 2004 Apr;6(4):497-509
PMID:16537923 - Sterol regulatory element binding protein is a principal regulator of anaerobic gene expression in fission yeast.
Todd BL et al. Mol Cell Biol 2006 Apr;26(7):2817-31
PMID:15659644 - Fission yeast mto2p regulates microtubule nucleation by the centrosomin-related protein mto1p.
Samejima I et al. Mol Biol Cell 2005 Jun;16(6):3040-51
PMID:23886939 - Functional replacement of fission yeast γ-tubulin small complex proteins Alp4 and Alp6 by human GCP2 and GCP3.
Riehlman TD et al. J Cell Sci 2013 Oct 01;126(Pt 19):4406-13
PMID:15004232 - Identification and characterization of two novel proteins affecting fission yeast gamma-tubulin complex function.
Venkatram S et al. Mol Biol Cell 2004 May;15(5):2287-301
PMID:20473289 - Analysis of a genome-wide set of gene deletions in the fission yeast Schizosaccharomyces pombe.
Kim DU et al. Nat Biotechnol 2010 Jun;28(6):617-623
PMID:35622906 - Distribution of γ-tubulin ring complex at the Schizosaccharomyces pombe spindle pole.
Saha S et al. MicroPubl Biol 2021;2021
PMID:39476757 - Characterization of Ksg1 protein kinase-dependent phosphoproteome in the fission yeast S. pombe.
Cipak L et al. Biochem Biophys Res Commun 2024 Oct 25;736:150895
PB_REF:0000006 - Disease associations from Monarch via human-pombe orthologs
PMID:15280226 - Functional dissection of the gamma-tubulin complex by suppressor analysis of gtb1 and alp4 mutations in Schizosaccharomyces pombe.
Tange Y et al. Genetics 2004 Jul;167(3):1095-107
PMID:16611237 - Modulation of Alp4 function in Schizosaccharomyces pombe induces novel phenotypes that imply distinct functions for nuclear and cytoplasmic gamma-tubulin complexes.
Masuda H et al. Genes Cells 2006 Apr;11(4):319-36
PMID:28619713 - Molecular model of fission yeast centrosome assembly determined by superresolution imaging.
Bestul AJ et al. J Cell Biol 2017 Aug 07;216(8):2409-2424
PMID:19001497 - Two distinct regions of Mto1 are required for normal microtubule nucleation and efficient association with the gamma-tubulin complex in vivo.
Samejima I et al. J Cell Sci 2008 Dec 01;121(Pt 23):3971-80
PMID:15837798 - Efficient formation of bipolar microtubule bundles requires microtubule-bound gamma-tubulin complexes.
Janson ME et al. J Cell Biol 2005 Apr 25;169(2):297-308
PMID:30427751 - The J-domain cochaperone Rsp1 interacts with Mto1 to organize noncentrosomal microtubule assembly.
Shen J et al. Mol Biol Cell 2019 Jan 15;30(2):256-267
PMID:35286199 - Mitotic spindle formation in the absence of Polo kinase.
Kim J et al. Proc Natl Acad Sci U S A 2022 Mar 22;119(12):e2114429119
GO_REF:0000033 - Annotation inferences using phylogenetic trees
PMID:31618856 - Two XMAP215/TOG Microtubule Polymerases, Alp14 and Dis1, Play Non-Exchangeable, Distinct Roles in Microtubule Organisation in Fission Yeast.
Yukawa M et al. Int J Mol Sci 2019 Oct 15;20(20)
PMID:23697806 - A genome-wide resource of cell cycle and cell shape genes of fission yeast.
Hayles J et al. Open Biol 2013 May 22;3(5):130053
PMID:16111942 - Hrs1p/Mcp6p on the meiotic SPB organizes astral microtubule arrays for oscillatory nuclear movement.
Tanaka K et al. Curr Biol 2005 Aug 23;15(16):1479-86
PMID:9658169 - Identification of novel temperature-sensitive lethal alleles in essential beta-tubulin and nonessential alpha 2-tubulin genes as fission yeast polarity mutants.
Radcliffe P et al. Mol Biol Cell 1998 Jul;9(7):1757-71
PMID:34346498 - Loss of kinesin-8 improves the robustness of the self-assembled spindle in Schizosaccharomyces pombe.
Pineda-Santaella A et al. J Cell Sci 2021 Aug 15;134(16)
PMID:20118936 - Schizosaccharomyces pombe genome-wide nucleosome mapping reveals positioning mechanisms distinct from those of Saccharomyces cerevisiae.
Lantermann AB et al. Nat Struct Mol Biol 2010 Feb;17(2):251-7
PMID:27889481 - Mitotic Nuclear Envelope Breakdown and Spindle Nucleation Are Controlled by Interphase Contacts between Centromeres and the Nuclear Envelope.
Fernández-Álvarez A et al. Dev Cell 2016 Dec 05;39(5):544-559
PMID:24006493 - Mzt1/Tam4, a fission yeast MOZART1 homologue, is an essential component of the γ-tubulin complex and directly interacts with GCP3(Alp6).
Dhani DK et al. Mol Biol Cell 2013 Nov;24(21):3337-49
PMID:11080156 - The fission yeast gamma-tubulin complex is required in G(1) phase and is a component of the spindle assembly checkpoint.
Vardy L et al. EMBO J 2000 Nov 15;19(22):6098-111
PMID:28017606 - Generation of a Spindle Checkpoint Arrest from Synthetic Signaling Assemblies.
Yuan I et al. Curr Biol 2017 Jan 09;27(1):137-143
PMID:29779879 - The XMAP215 Ortholog Alp14 Promotes Microtubule Nucleation in Fission Yeast.
Flor-Parra I et al. Curr Biol 2018 Jun 04;28(11):1681-1691.e4
PMID:16855399 - An extended anaphase signaling pathway for Mad2p includes microtubule organizing center proteins and multiple motor-dependent transitions.
Mayer C et al. Cell Cycle 2006 Jul;5(13):1456-63
PMID:19942852 - Fission yeast Pcp1 links polo kinase-mediated mitotic entry to gamma-tubulin-dependent spindle formation.
Fong CS et al. EMBO J 2010 Jan 06;29(1):120-30
PMID:30726745 - Fission Yeast NDR/LATS Kinase Orb6 Regulates Exocytosis via Phosphorylation of the Exocyst Complex.
Tay YD et al. Cell Rep 2019 Feb 05;26(6):1654-1667.e7
PMID:24943839 - A role for nuclear envelope-bridging complexes in homology-directed repair.
Swartz RK et al. Mol Biol Cell 2014 Aug 15;25(16):2461-71
PMID:12134075 - A fourth component of the fission yeast gamma-tubulin complex, Alp16, is required for cytoplasmic microtubule integrity and becomes indispensable when gamma-tubulin function is compromised.
Fujita A et al. Mol Biol Cell 2002 Jul;13(7):2360-73
PMID:33823663 - A TOR (target of rapamycin) and nutritional phosphoproteome of fission yeast reveals novel targets in networks conserved in humans.
Halova L et al. Open Biol 2021 Apr;11(4):200405
PMID:15772152 - Effects of {gamma}-tubulin complex proteins on microtubule nucleation and catastrophe in fission yeast.
Zimmerman S et al. Mol Biol Cell 2005 Jun;16(6):2719-33
PMID:17486116 - Gamma-tubulin complex-mediated anchoring of spindle microtubules to spindle-pole bodies requires Msd1 in fission yeast.
Toya M et al. Nat Cell Biol 2007 Jun;9(6):646-53
PMID:21993292 - Microtubule nucleation by γ-tubulin complexes.
Kollman JM et al. Nat Rev Mol Cell Biol 2011 Oct 12;12(11):709-21
PMID:15120067 - Microtubule nucleation at non-spindle pole body microtubule-organizing centers requires fission yeast centrosomin-related protein mod20p.
Sawin KE et al. Curr Biol 2004 May 04;14(9):763-75
PMID:16611238 - The carboxy-terminus of Alp4 alters microtubule dynamics to induce oscillatory nuclear movement led by the spindle pole body in Schizosaccharomyces pombe.
Masuda H et al. Genes Cells 2006 Apr;11(4):337-52
PMID:23101633 - Quantitative analysis of fission yeast transcriptomes and proteomes in proliferating and quiescent cells.
Marguerat S et al. Cell 2012 Oct 26;151(3):671-83
PMID:11952833 - The gamma-tubulin complex protein Alp4 provides a link between the metaphase checkpoint and cytokinesis in fission yeast.
Vardy L et al. Genes Cells 2002 Apr;7(4):365-73
PMID:20833892 - Schizosaccharomyces pombe calmodulin, Cam1, plays a crucial role in sporulation by recruiting and stabilizing the spindle pole body components responsible for assembly of the forespore membrane.
Itadani A et al. Eukaryot Cell 2010 Dec;9(12):1925-35
PMID:23885124 - Fission yeast MOZART1/Mzt1 is an essential γ-tubulin complex component required for complex recruitment to the microtubule organizing center, but not its assembly.
Masuda H et al. Mol Biol Cell 2013 Sep;24(18):2894-906
PMID:25720772 - Quantitative phosphoproteomics reveals pathways for coordination of cell growth and division by the conserved fission yeast kinase pom1.
Kettenbach AN et al. Mol Cell Proteomics 2015 May;14(5):1275-87
PMID:24763107 - Absolute proteome and phosphoproteome dynamics during the cell cycle of Schizosaccharomyces pombe (Fission Yeast).
Carpy A et al. Mol Cell Proteomics 2014 Aug;13(8):1925-36
PMID:17021256 - Noncore components of the fission yeast gamma-tubulin complex.
Anders A et al. Mol Biol Cell 2006 Dec;17(12):5075-93
PMID:21511999 - Comparative functional genomics of the fission yeasts.
Rhind N et al. Science 2011 May 20;332(6032):930-6
PMID:23388459 - Kinesin-14 Pkl1 targets γ-tubulin for release from the γ-tubulin ring complex (γ-TuRC) ‬‬‬‬‬‬‬.
Olmsted ZT et al. Cell Cycle 2013 Mar 01;12(5):842-8
PMID:25987607 - The Msd1-Wdr8-Pkl1 complex anchors microtubule minus ends to fission yeast spindle pole bodies.
Yukawa M et al. J Cell Biol 2015 May 25;209(4):549-62
PMID:15569158 - The Sec14 family glycerophospholipid-transfer protein is required for structural integrity of the spindle pole body during meiosis in fission yeast.
Nakase Y et al. Genes Cells 2004 Dec;9(12):1275-86
PMID:26771498 - A Proteome-wide Fission Yeast Interactome Reveals Network Evolution Principles from Yeasts to Human.
Vo TV et al. Cell 2016 Jan 14;164(1-2):310-323
PMID:25103238 - Tts1, the fission yeast homologue of the TMEM33 family, functions in NE remodeling during mitosis.
Zhang D et al. Mol Biol Cell 2014 Oct 01;25(19):2970-83
PMID:22425159 - Transient structure associated with the spindle pole body directs meiotic microtubule reorganization in S. pombe.
Funaya C et al. Curr Biol 2012 Apr 10;22(7):562-74
PMID:18418055 - Protein complexes at the microtubule organizing center regulate bipolar spindle assembly.
Rodriguez AS et al. Cell Cycle 2008 May 01;7(9):1246-53
PMID:26537787 - Targeting of SUMO substrates to a Cdc48-Ufd1-Npl4 segregase and STUbL pathway in fission yeast.
Køhler JB et al. Nat Commun 2015 Nov 05;6:8827
PMID:39367033 - Quantitative proteomics and phosphoproteomics profiling of meiotic divisions in the fission yeast Schizosaccharomyces pombe.
Sivakova B et al. Sci Rep 2024 Oct 04;14(1):23105
PMID:12161753 - The transcriptional program of meiosis and sporulation in fission yeast.
Mata J et al. Nat Genet 2002 Sep;32(1):143-7
PMID:18257517 - Phosphoproteome analysis of fission yeast.
Wilson-Grady JT et al. J Proteome Res 2008 Mar;7(3):1088-97
PMID:21340088 - Microarray-based target identification using drug hypersensitive fission yeast expressing ORFeome.
Arita Y et al. Mol Biosyst 2011 May;7(5):1463-72
PMID:29996109 - Quantitative Phosphoproteomics Reveals the Signaling Dynamics of Cell-Cycle Kinases in the Fission Yeast Schizosaccharomyces pombe.
Swaffer MP et al. Cell Rep 2018 Jul 10;24(2):503-514
PMID:21444751 - Microtubule stabilization in vivo by nucleation-incompetent γ-tubulin complex.
Anders A et al. J Cell Sci 2011 Apr 15;124(Pt 8):1207-13
PMID:21712547 - Mitotic substrates of the kinase aurora with roles in chromatin regulation identified through quantitative phosphoproteomics of fission yeast.
Koch A et al. Sci Signal 2011 Jun 28;4(179):rs6
PMID:16823372 - ORFeome cloning and global analysis of protein localization in the fission yeast Schizosaccharomyces pombe.
Matsuyama A et al. Nat Biotechnol 2006 Jul;24(7):841-7
PMID:37783794 - An ESCRT grommet cooperates with a diffusion barrier to maintain nuclear integrity.
Ader NR et al. Nat Cell Biol 2023 Oct;25(10):1465-1477
PMID:15800064 - Mto2p, a novel fission yeast protein required for cytoplasmic microtubule organization and anchoring of the cytokinetic actin ring.
Venkatram S et al. Mol Biol Cell 2005 Jun;16(6):3052-63
PMID:25348260 - Kinesin-14 and kinesin-5 antagonistically regulate microtubule nucleation by γ-TuRC in yeast and human cells.
Olmsted ZT et al. Nat Commun 2014 Oct 28;5:5339
PMID:27053664 - Synergistic role of fission yeast Alp16GCP6 and Mzt1MOZART1 in γ-tubulin complex recruitment to mitotic spindle pole bodies and spindle assembly.
Masuda H et al. Mol Biol Cell 2016 Jun 01;27(11):1753-63