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protein coding gene - mad1 (SPBC3D6.04c) - Mad2 receptor at unattached kinetochores, Mad1

Gene summary

Standard name
mad1
Systematic ID
SPBC3D6.04c
Product
Mad2 receptor at unattached kinetochores, Mad1
Organism
Schizosaccharomyces pombe (fission yeast)
UniProt ID
P87169
ORFeome ID
37/37D06
Characterisation status
biological role published
Feature type
mRNA gene
Genomic location
chromosome II: 1274839..1277073 reverse strand

Annotation

Comment

PBO:0109299 - coding sequence mRNA is required to establish mad1+-like expression

References:

PBO:0109297 - codon-optimization decreases protein concentration

References:

PBO:0109298 - Mad1 homodimers assemble co-translationally

References:

Disease association

MONDO:0859346 - mosaic variegated aneuploidy syndrome 7 with inflammation and tumor predisposition

References:

MONDO:0700275 - prostate cancer, hereditary

References:

GO biological process

GO:0051315 - attachment of mitotic spindle microtubules to kinetochore

References:

GO:0099606 - microtubule plus-end directed mitotic chromosome migration

References:

GO:0007094 - mitotic spindle assembly checkpoint signaling

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GO cellular component

GO:0000776 - kinetochore

References:

GO:0072686 - mitotic spindle

References:

GO:0044732 - mitotic spindle pole body

References:

GO:0005635 - nuclear envelope

References:

GO:0005634 - nucleus

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GO molecular function

GO:0140483 - kinetochore adaptor activity

References:

GO:0005515 - protein binding

References:

Modification

MOD:00046 - O-phospho-L-serine

References:

MOD:00047 - O-phospho-L-threonine

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Multi-locus phenotype

FYPO:0004318 - abolished mitotic spindle assembly checkpoint

References:

Genotypes:

FYPO:0000941 - abolished protein localization to mitotic spindle pole body

References:

Genotypes:

FYPO:0000082 - decreased cell population growth at high temperature

References:

Genotypes:

FYPO:0002638 - increased activation of mitotic spindle assembly checkpoint

References:

Genotypes:

FYPO:0005684 - increased duration of mitotic prometaphase

References:

Genotypes:

FYPO:0002061 - inviable vegetative cell population

References:

Genotypes:

FYPO:0000324 - mitotic metaphase/anaphase transition delay

References:

Genotypes:

FYPO:0003762 - normal mitotic spindle assembly checkpoint

References:

Genotypes:

FYPO:0003241 - unequal mitotic sister chromatid segregation

References:

Genotypes:

FYPO:0002060 - viable vegetative cell population

References:

Genotypes:

Quantitative gene expression

PBO:0006310 - protein level

References:

PBO:0011963 - RNA level

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Single locus phenotype

FYPO:0000059 - abnormal mitotic cell cycle

References:

Genotypes:

FYPO:0000030 - abnormal mitotic chromosome congression

References:

Genotypes:

FYPO:0008164 - abnormal protein localization to kinetochore during mitotic M phase

References:

Genotypes:

FYPO:0004318 - abolished mitotic spindle assembly checkpoint

References:

Genotypes:

FYPO:0007383 - abolished protein localization to kinetochore during mitotic spindle assembly checkpoint signaling

References:

Genotypes:

FYPO:0002568 - abolished protein localization to nuclear periphery

References:

Genotypes:

FYPO:0000705 - abolished protein-protein interaction

References:

Genotypes:

FYPO:0005781 - decreased duration of mitotic spindle assembly checkpoint

References:

Genotypes:

FYPO:0001324 - decreased protein level during vegetative growth

References:

Genotypes:

FYPO:0005300 - decreased protein localization to kinetochore during mitotic spindle assembly checkpoint signaling

References:

Genotypes:

FYPO:0005211 - decreased protein localization to mitotic spindle pole body during anaphase

References:

Genotypes:

FYPO:0001645 - decreased protein-protein interaction

References:

Genotypes:

FYPO:0001122 - elongated vegetative cell

References:

Genotypes:

FYPO:0003938 - increased cell population growth during glucose starvation

References:

Genotypes:

FYPO:0000636 - increased cell population growth rate

References:

Genotypes:

FYPO:0001327 - increased protein level during vegetative growth

References:

Genotypes:

FYPO:0001309 - increased viability in stationary phase

References:

Genotypes:

FYPO:0006518 - loss of viability in G0

References:

Genotypes:

FYPO:0001383 - normal DNA content

References:

Genotypes:

FYPO:0007553 - normal G1 to G0 transition

References:

Genotypes:

FYPO:0003762 - normal mitotic spindle assembly checkpoint

References:

Genotypes:

FYPO:0005042 - normal protein localization to kinetochore

References:

Genotypes:

FYPO:0006641 - normal protein localization to kinetochore during mitotic metaphase

References:

Genotypes:

FYPO:0005212 - normal protein localization to kinetochore during mitotic prometaphase

References:

Genotypes:

FYPO:0002901 - normal protein localization to kinetochore during vegetative growth

References:

Genotypes:

FYPO:0002563 - normal protein localization to nuclear periphery

References:

Genotypes:

FYPO:0000703 - normal protein-protein interaction

References:

Genotypes:

FYPO:0000067 - resistance to brefeldin A

References:

Genotypes:

FYPO:0009079 - resistance to calcofluor and sodium dodecyl sulfate

References:

Genotypes:

FYPO:0002693 - resistance to diamide

References:

Genotypes:

FYPO:0009043 - resistance to potassium chloride and sodium dodecyl sulfate

References:

Genotypes:

FYPO:0004432 - sensitive to 2-bromo-1-(4-methoxyphenyl)-3-((4-methylphenyl)sulfonyl)-1-propanone

References:

Genotypes:

FYPO:0006680 - sensitive to bisphenol A

References:

Genotypes:

FYPO:0000095 - sensitive to bleomycin

References:

Genotypes:

FYPO:0003840 - sensitive to carbendazim

References:

Genotypes:

FYPO:0007931 - sensitive to egtazic acid

References:

Genotypes:

FYPO:0000842 - sensitive to ethanol during vegetative growth

References:

Genotypes:

FYPO:0000785 - sensitive to formamide

References:

Genotypes:

FYPO:0000088 - sensitive to hydroxyurea

References:

Genotypes:

FYPO:0000091 - sensitive to thiabendazole

References:

Genotypes:

FYPO:0000268 - sensitive to UV during vegetative growth

References:

Genotypes:

FYPO:0004433 - sensitive to viridicatumtoxin

References:

Genotypes:

FYPO:0005069 - unequal mitotic sister chromatid segregation following normal mitosis

References:

Genotypes:

FYPO:0002060 - viable vegetative cell population

References:

Genotypes:

FYPO:0002104 - viable vegetative cell with normal cell shape

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)

Warnings

PBO:0000070 - gene structure updated

References:

Protein features

IDNameInterPro nameDB name
PF05557MADMad1PFAM
G3DSA:1.20.5.170GENE3D
G3DSA:3.30.457.60GENE3D
PTHR23168MITOTIC SPINDLE ASSEMBLY CHECKPOINT PROTEIN MAD1 MITOTIC ARREST DEFICIENT-LIKE PROTEIN 1Mad1PANTHER
CoilCoilCOILS
mobidb-lite-Disorderdisorder_predictionMOBIDB-Disorder

Orthologs

References / Literature

PMID:16394105 - The V260I mutation in fission yeast alpha-tubulin Atb2 affects microtubule dynamics and EB1-Mal3 localization and activates the Bub1 branch of the spindle checkpoint.
Asakawa K et al. Mol Biol Cell 2006 Mar;17(3):1421-35
PMID:15483052 - BAF53/Arp4 homolog Alp5 in fission yeast is required for histone H4 acetylation, kinetochore-spindle attachment, and gene silencing at centromere.
Minoda A et al. Mol Biol Cell 2005 Jan;16(1):316-27
PMID:12951601 - Deletion of Mia1/Alp7 activates Mad2-dependent spindle assembly checkpoint in fission yeast.
Sato M et al. Nat Cell Biol 2003 Sep;5(9):764-6; author reply 766
PB_REF:0000006 - Disease associations from Monarch via human-pombe orthologs
PMID:11950879 - Control of localization of a spindle checkpoint protein, Mad2, in fission yeast.
Ikui AE et al. J Cell Sci 2002 Apr 15;115(Pt 8):1603-10
PMID:18204818 - Identification of small molecules inducing apoptosis by cell-based assay using fission yeast deletion mutants.
Chung KS et al. Invest New Drugs 2008 Aug;26(4):299-307
PMID:22184248 - Centromere-tethered Mps1 pombe homolog (Mph1) kinase is a sufficient marker for recruitment of the spindle checkpoint protein Bub1, but not Mad1.
Ito D et al. Proc Natl Acad Sci U S A 2012 Jan 03;109(1):209-14
PMID:28366743 - Different Functionality of Cdc20 Binding Sites within the Mitotic Checkpoint Complex.
Sewart K et al. Curr Biol 2017 Apr 24;27(8):1213-1220
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:26258632 - Mad1 promotes chromosome congression by anchoring a kinesin motor to the kinetochore.
Akera T et al. Nat Cell Biol 2015 Sep;17(9):1124-33
PMID:32435206 - Posttranslational Arginylation Enzyme Arginyltransferase1 Shows Genetic Interactions With Specific Cellular Pathways in vivo .
Wiley DJ et al. Front Physiol 2020;11:427
PMID:33260998 - High-Throughput Flow Cytometry Combined with Genetic Analysis Brings New Insights into the Understanding of Chromatin Regulation of Cellular Quiescence.
Zahedi Y et al. Int J Mol Sci 2020 Nov 27;21(23)
PMID:38166399 - Fission yeast Wee1 is required for stable kinetochore-microtubule attachment.
Takado M et al. Open Biol 2024 Jan;14(1):230379
PMID:18818364 - Conservation and rewiring of functional modules revealed by an epistasis map in fission yeast.
Roguev A et al. Science 2008 Oct 17;322(5900):405-10
PMID:28357272 - A central role for TOR signalling in a yeast model for juvenile CLN3 disease.
Bond ME et al. Microb Cell 2015 Nov 11;2(12):466-480
PMID:19523829 - Phosphorylation state defines discrete roles for monopolin in chromosome attachment and spindle elongation.
Choi SH et al. Curr Biol 2009 Jun 23;19(12):985-95
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:26900649 - The hairpin region of Ndc80 is important for the kinetochore recruitment of Mph1/MPS1 in fission yeast.
Chmielewska AE et al. Cell Cycle 2016;15(5):740-7
PMID:18562692 - Latrunculin A delays anaphase onset in fission yeast by disrupting an Ase1-independent pathway controlling mitotic spindle stability.
Meadows JC et al. Mol Biol Cell 2008 Sep;19(9):3713-23
PMID:28178520 - Identification of a Sgo2-Dependent but Mad2-Independent Pathway Controlling Anaphase Onset in Fission Yeast.
Meadows JC et al. Cell Rep 2017 Feb 07;18(6):1422-1433
PMID:22681890 - Hierarchical modularity and the evolution of genetic interactomes across species.
Ryan CJ et al. Mol Cell 2012 Jun 08;46(5):691-704
GO_REF:0000033 - Annotation inferences using phylogenetic trees
PMID:23163955 - Analysis of stress-induced duplex destabilization (SIDD) properties of replication origins, genes and intergenes in the fission yeast, Schizosaccharomyces pombe.
Yadav MP et al. BMC Res Notes 2012 Nov 19;5:643
PMID:22825872 - Mph1 kinetochore localization is crucial and upstream in the hierarchy of spindle assembly checkpoint protein recruitment to kinetochores.
Heinrich S et al. J Cell Sci 2012 Oct 15;125(Pt 20):4720-7
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:20537132 - Global fitness profiling of fission yeast deletion strains by barcode sequencing.
Han TX et al. Genome Biol 2010;11(6):R60
PMID:11909965 - Fission yeast Mad3p is required for Mad2p to inhibit the anaphase-promoting complex and localizes to kinetochores in a Bub1p-, Bub3p-, and Mph1p-dependent manner.
Millband DN et al. Mol Cell Biol 2002 Apr;22(8):2728-42
PMID:24583014 - Slow checkpoint activation kinetics as a safety device in anaphase.
Kamenz J et al. Curr Biol 2014 Mar 17;24(6):646-51
PMID:18931302 - Significant conservation of synthetic lethal genetic interaction networks between distantly related eukaryotes.
Dixon SJ et al. Proc Natl Acad Sci U S A 2008 Oct 28;105(43):16653-8
PMID:23166349 - Csi1 links centromeres to the nuclear envelope for centromere clustering.
Hou H et al. J Cell Biol 2012 Nov 26;199(5):735-44
PMID:24161933 - Determinants of robustness in spindle assembly checkpoint signalling.
Heinrich S et al. Nat Cell Biol 2013 Nov;15(11):1328-39
PMID:22281223 - Kinase activity of fission yeast Mph1 is required for Mad2 and Mad3 to stably bind the anaphase promoting complex.
Zich J et al. Curr Biol 2012 Feb 21;22(4):296-301
PMID:30647105 - Comparative Genomic Screen in Two Yeasts Reveals Conserved Pathways in the Response Network to Phenol Stress.
Alhoch B et al. G3 (Bethesda) 2019 Mar 07;9(3):639-650
PMID:28974540 - The fission yeast nucleoporin Alm1 is required for proteasomal degradation of kinetochore components.
Salas-Pino S et al. J Cell Biol 2017 Nov 06;216(11):3591-3608
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:24477934 - Mad1 contribution to spindle assembly checkpoint signalling goes beyond presenting Mad2 at kinetochores.
Heinrich S et al. EMBO Rep 2014 Mar;15(3):291-8
PMID:22521786 - Phosphodependent recruitment of Bub1 and Bub3 to Spc7/KNL1 by Mph1 kinase maintains the spindle checkpoint.
Shepperd LA et al. Curr Biol 2012 May 22;22(10):891-9
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:25452419 - Parallel profiling of fission yeast deletion mutants for proliferation and for lifespan during long-term quiescence.
Sideri T et al. G3 (Bethesda) 2014 Dec 01;5(1):145-55
PMID:19680287 - Bub1 and Bub3 promote the conversion from monopolar to bipolar chromosome attachment independently of shugoshin.
Windecker H et al. EMBO Rep 2009 Sep;10(9):1022-8
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:30355493 - Expanded Interactome of the Intrinsically Disordered Protein Dss1.
Schenstrøm SM et al. Cell Rep 2018 Oct 23;25(4):862-870
PMID:35811551 - Mitotic checkpoint gene expression is tuned by codon usage bias.
Esposito E et al. EMBO J 2022 Aug 01;41(15):e107896
PMID:35820914 - Antagonistic effects of mitochondrial matrix and intermembrane space proteases on yeast aging.
Vega M et al. BMC Biol 2022 Jul 12;20(1):160
PMID:23173672 - Identification of novel genes involved in DNA damage response by screening a genome-wide Schizosaccharomyces pombe deletion library.
Pan X et al. BMC Genomics 2012 Nov 23;13:662
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:22660415 - MPS1/Mph1 phosphorylates the kinetochore protein KNL1/Spc7 to recruit SAC components.
Yamagishi Y et al. Nat Cell Biol 2012 Jun 03;14(7):746-52
PMID:16360688 - Control of Shugoshin function during fission-yeast meiosis.
Vaur S et al. Curr Biol 2005 Dec 20;15(24):2263-70
PMID:21511999 - Comparative functional genomics of the fission yeasts.
Rhind N et al. Science 2011 May 20;332(6032):930-6
PMID:22437499 - Structure of the mitotic checkpoint complex.
Chao WC et al. Nature 2012 Mar 21;484(7393):208-13
PMID:29021344 - A microtubule polymerase cooperates with the kinesin-6 motor and a microtubule cross-linker to promote bipolar spindle assembly in the absence of kinesin-5 and kinesin-14 in fission yeast.
Yukawa M et al. Mol Biol Cell 2017 Dec 01;28(25):3647-3659
PMID:31257143 - The Bub1-TPR Domain Interacts Directly with Mad3 to Generate Robust Spindle Checkpoint Arrest.
Leontiou I et al. Curr Biol 2019 Jul 22;29(14):2407-2414.e7
PMID:15347659 - DNA replication checkpoint control mediated by the spindle checkpoint protein Mad2p in fission yeast.
Sugimoto I et al. J Biol Chem 2004 Nov 05;279(45):47372-8
PMID:27618268 - Bub3-Bub1 Binding to Spc7/KNL1 Toggles the Spindle Checkpoint Switch by Licensing the Interaction of Bub1 with Mad1-Mad2.
Mora-Santos MD et al. Curr Biol 2016 Oct 10;26(19):2642-2650
PMID:37787768 - Broad functional profiling of fission yeast proteins using phenomics and machine learning.
Rodríguez-López M et al. Elife 2023 Oct 03;12