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protein coding gene - hsk1 (SPBC776.12c) - Dbf4(Dfp1)-dependent protein kinase Hsk1

Gene summary

Standard name
hsk1
Systematic ID
SPBC776.12c
Product
Dbf4(Dfp1)-dependent protein kinase Hsk1
Organism
Schizosaccharomyces pombe (fission yeast)
Synonyms
cdc7
UniProt ID
P50582
ORFeome ID
40/40F05
Characterisation status
biological role published
Feature type
mRNA gene
Genomic location
chromosome II: 3197060..3199201 reverse strand

Annotation

GO biological process

GO:0006270 - DNA replication initiation

References:

GO:0000727 - double-strand break repair via break-induced replication

References:

GO:0000082 - G1/S transition of mitotic cell cycle

References:

GO:0033314 - mitotic DNA replication checkpoint signaling

References:

GO:0031573 - mitotic intra-S DNA damage checkpoint signaling

References:

GO:0045739 - positive regulation of DNA repair

References:

GO:1903468 - positive regulation of DNA replication initiation

References:

GO cellular component

GO:0005737 - cytoplasm

References:

GO:0031431 - Dbf4-dependent protein kinase complex

References:

GO:0005634 - nucleus

References:

GO molecular function

GO:0005524 - ATP binding

References:

GO:0004674 - protein serine/threonine kinase activity

References:

Modification

MOD:00046 - O-phospho-L-serine

References:

MOD:00047 - O-phospho-L-threonine

References:

MOD:00696 - phosphorylated residue

References:

Multi-locus phenotype

FYPO:0000848 - abnormal chromosome morphology during vegetative growth

References:

Genotypes:

FYPO:0000444 - abnormal mitotic cell cycle arrest with replicated DNA

References:

Genotypes:

FYPO:0000141 - abnormal mitotic sister chromatid segregation

References:

Genotypes:

FYPO:0003263 - abnormal sporulation resulting in formation of ascus with more than four spores

References:

Genotypes:

FYPO:0002196 - abnormal vegetative cell shape

References:

Genotypes:

FYPO:0004918 - abolished positive regulation of protein kinase activity

References:

Genotypes:

FYPO:0003165 - cut with abnormal chromosome segregation

References:

Genotypes:

FYPO:0001250 - decreased origin firing efficiency

References:

Genotypes:

FYPO:0001382 - decreased protein kinase activity

References:

Genotypes:

FYPO:0002098 - decreased protein phosphorylation during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0001645 - decreased protein-protein interaction

References:

Genotypes:

FYPO:0000584 - decreased sporulation frequency

References:

Genotypes:

FYPO:0001355 - decreased vegetative cell population growth

References:

Genotypes:

FYPO:0000614 - increased duration of mitotic S phase

References:

Genotypes:

FYPO:0001249 - increased origin firing efficiency

References:

Genotypes:

FYPO:0002680 - increased protein phosphorylation

References:

Genotypes:

FYPO:0000839 - inviable elongated mononucleate aseptate cell

References:

Genotypes:

FYPO:0004672 - inviable elongated vegetative cell with fragmented nucleus

References:

Genotypes:

FYPO:0002061 - inviable vegetative cell population

References:

Genotypes:

FYPO:0003082 - normal genetic imprinting at mating-type locus

References:

Genotypes:

FYPO:0000963 - normal growth on hydroxyurea

References:

Genotypes:

FYPO:0001315 - normal vegetative cell morphology

References:

Genotypes:

FYPO:0001357 - normal vegetative cell population growth

References:

Genotypes:

FYPO:0000416 - premature mitotic sister chromatid separation

References:

Genotypes:

FYPO:0000085 - sensitive to camptothecin

References:

Genotypes:

FYPO:0000088 - sensitive to hydroxyurea

References:

Genotypes:

FYPO:0000089 - sensitive to methyl methanesulfonate

References:

Genotypes:

FYPO:0000268 - sensitive to UV during vegetative growth

References:

Genotypes:

FYPO:0003241 - unequal mitotic sister chromatid segregation

References:

Genotypes:

FYPO:0002060 - viable vegetative cell population

References:

Genotypes:

Qualitative gene expression

PomGeneEx:0000018 - protein level increased

References:

PomGeneEx:0000012 - RNA level decreased

References:

PomGeneEx:0000011 - RNA level increased

References:

Quantitative gene expression

PBO:0006310 - protein level

References:

PBO:0011963 - RNA level

References:

Single locus phenotype

FYPO:0000848 - abnormal chromosome morphology during vegetative growth

References:

Genotypes:

FYPO:0000059 - abnormal mitotic cell cycle

References:

Genotypes:

FYPO:0000444 - abnormal mitotic cell cycle arrest with replicated DNA

References:

Genotypes:

FYPO:0001430 - abnormal mitotic cell cycle arrest with unreplicated DNA

References:

Genotypes:

FYPO:0001933 - abnormal mitotic cell cycle regulation during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0001343 - abnormal mitotic DNA replication

References:

Genotypes:

FYPO:0000141 - abnormal mitotic sister chromatid segregation

References:

Genotypes:

FYPO:0000623 - abnormal positive regulation of transcription

References:

Genotypes:

FYPO:0003019 - abnormal protein autophosphorylation during vegetative growth

References:

Genotypes:

FYPO:0002196 - abnormal vegetative cell shape

References:

Genotypes:

FYPO:0001093 - abolished chromatin binding

References:

Genotypes:

FYPO:0004918 - abolished positive regulation of protein kinase activity

References:

Genotypes:

FYPO:0001384 - abolished protein kinase activity

References:

Genotypes:

FYPO:0005095 - abolished protein localization to chromatin at replication origin

References:

Genotypes:

FYPO:0002779 - abolished protein localization to nucleus, with protein mislocalized to cytoplasm, during vegetative growth

References:

Genotypes:

FYPO:0004550 - abolished protein phosphorylation during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0004660 - abolished protein phosphorylation during cellular response to methyl methanesulfonate

References:

Genotypes:

FYPO:0000229 - cut

References:

Genotypes:

FYPO:0003165 - cut with abnormal chromosome segregation

References:

Genotypes:

FYPO:0005093 - decreased duration of protein phosphorylation during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0005094 - decreased duration of S-phase DNA damage checkpoint during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0004240 - decreased histone H3-K56 acetylation during vegetative growth

References:

Genotypes:

FYPO:0001250 - decreased origin firing efficiency

References:

Genotypes:

FYPO:0007878 - decreased protein degradation during cellular response to methyl methanesulfonate

References:

Genotypes:

FYPO:0001382 - decreased protein kinase activity

References:

Genotypes:

FYPO:0002097 - decreased protein kinase activity during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0003950 - decreased protein localization to chromatin at replication origin

References:

Genotypes:

FYPO:0002909 - decreased protein localization to chromatin during vegetative growth

References:

Genotypes:

FYPO:0004056 - decreased protein localization to nucleus, with protein mislocalized to cytoplasm

References:

Genotypes:

FYPO:0002098 - decreased protein phosphorylation during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0002470 - decreased protein phosphorylation during cellular response to ionizing radiation

References:

Genotypes:

FYPO:0001645 - decreased protein-protein interaction

References:

Genotypes:

FYPO:0001355 - decreased vegetative cell population growth

References:

Genotypes:

FYPO:0000453 - DNA content decreased during vegetative growth

References:

Genotypes:

FYPO:0001386 - increased haploidization

References:

Genotypes:

FYPO:0001974 - increased number of cells with 1C DNA content

References:

Genotypes:

FYPO:0000786 - increased plasmid loss

References:

Genotypes:

FYPO:0000847 - increased protein degradation during vegetative growth

References:

Genotypes:

FYPO:0001327 - increased protein level during vegetative growth

References:

Genotypes:

FYPO:0001038 - increased protein phosphorylation during vegetative growth

References:

Genotypes:

FYPO:0000316 - inviable after spore germination

References:

Genotypes:

FYPO:0000314 - inviable after spore germination with elongated germ tube

References:

Genotypes:

FYPO:0001042 - inviable after spore germination, single or double cell division

References:

Genotypes:

FYPO:0000839 - inviable elongated mononucleate aseptate cell

References:

Genotypes:

FYPO:0004672 - inviable elongated vegetative cell with fragmented nucleus

References:

Genotypes:

FYPO:0002061 - inviable vegetative cell population

References:

Genotypes:

FYPO:0001387 - loss of viability at high temperature

References:

Genotypes:

FYPO:0000333 - mitotic G1/S phase transition delay

References:

Genotypes:

FYPO:0001383 - normal DNA content

References:

Genotypes:

FYPO:0004921 - normal positive regulation of protein kinase activity

References:

Genotypes:

FYPO:0002800 - normal protein degradation during vegetative growth

References:

Genotypes:

FYPO:0003075 - normal protein kinase activity

References:

Genotypes:

FYPO:0004548 - normal protein kinase activity during cellular response to hydroxyurea

References:

Genotypes:

FYPO:0000833 - normal protein level during vegetative growth

References:

Genotypes:

FYPO:0004962 - normal protein localization to chromatin at replication origin

References:

Genotypes:

FYPO:0001509 - normal protein localization to chromatin during vegetative growth

References:

Genotypes:

FYPO:0000838 - normal protein localization to nucleus during vegetative growth

References:

Genotypes:

FYPO:0000703 - normal protein-protein interaction

References:

Genotypes:

FYPO:0001315 - normal vegetative cell morphology

References:

Genotypes:

FYPO:0001357 - normal vegetative cell population growth

References:

Genotypes:

FYPO:0000416 - premature mitotic sister chromatid separation

References:

Genotypes:

FYPO:0000085 - sensitive to camptothecin

References:

Genotypes:

FYPO:0000088 - sensitive to hydroxyurea

References:

Genotypes:

FYPO:0000089 - sensitive to methyl methanesulfonate

References:

Genotypes:

FYPO:0000091 - sensitive to thiabendazole

References:

Genotypes:

FYPO:0000268 - sensitive to UV during vegetative growth

References:

Genotypes:

FYPO:0003241 - unequal mitotic sister chromatid segregation

References:

Genotypes:

FYPO:0002060 - viable vegetative cell population

References:

Genotypes:

Subunit composition

PBO:0015221 - heteromeric(2)

References:

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
PF00069PkinaseProt_kinase_domPFAM
cd14019STKc_Cdc7CDD
PS00108PROTEIN_KINASE_STSer/Thr_kinase_ASPROSITE_PATTERNS
PS50011PROTEIN_KINASE_DOMProt_kinase_domPROSITE_PROFILES
SM00220serkin_6Prot_kinase_domSMART
SSF56112Protein kinase-like (PK-like)Kinase-like_dom_sfSUPERFAMILY
G3DSA:3.30.200.20Phosphorylase Kinase; domain 1GENE3D
G3DSA:1.10.510.10Transferase(Phosphotransferase) domain 1GENE3D
PTHR44167OVARIAN-SPECIFIC SERINE/THREONINE-PROTEIN KINASE LOK-RELATEDPANTHER
mobidb-lite-Disorderdisorder_predictionMOBIDB-Disorder

Orthologs

References / Literature

PMID:21099360 - Hsk1 kinase and Cdc45 regulate replication stress-induced checkpoint responses in fission yeast.
Matsumoto S et al. Cell Cycle 2010 Dec 01;9(23):4627-37
PMID:10545452 - Drc1p/Cps1p, a 1,3-beta-glucan synthase subunit, is essential for division septum assembly in Schizosaccharomyces pombe.
Liu J et al. Genetics 1999 Nov;153(3):1193-203
PMID:22279046 - Rif1 is a global regulator of timing of replication origin firing in fission yeast.
Hayano M et al. Genes Dev 2012 Jan 15;26(2):137-50
PMID:23045396 - Continued DNA synthesis in replication checkpoint mutants leads to fork collapse.
Sabatinos SA et al. Mol Cell Biol 2012 Dec;32(24):4986-97
PMID:9705352 - Purification of Hsk1, a minichromosome maintenance protein kinase from fission yeast.
Brown GW et al. J Biol Chem 1998 Aug 21;273(34):22083-90
PMID:20230746 - Hsk1- and SCF(Pof3)-dependent proteolysis of S. pombe Ams2 ensures histone homeostasis and centromere function.
Takayama Y et al. Dev Cell 2010 Mar 16;18(3):385-96
PMID:11359920 - Regulation of initiation of S phase, replication checkpoint signaling, and maintenance of mitotic chromosome structures during S phase by Hsk1 kinase in the fission yeast.
Takeda T et al. Mol Biol Cell 2001 May;12(5):1257-74
PMID:14508607 - Ibp1p, a novel Cdc25-related phosphatase, suppresses Schizosaccharomyces pombe hsk1 ( cdc7).
Snaith HA et al. Curr Genet 2003 Oct;44(1):38-48
PMID:12972571 - Fission yeast Cdc23/Mcm10 functions after pre-replicative complex formation to promote Cdc45 chromatin binding.
Gregan J et al. Mol Biol Cell 2003 Sep;14(9):3876-87
PMID:11027263 - Schizosaccharomyces pombe Hsk1p is a potential cds1p target required for genome integrity.
Snaith HA et al. Mol Cell Biol 2000 Nov;20(21):7922-32
PMID:11402029 - Bipartite binding of a kinase activator activates Cdc7-related kinase essential for S phase.
Ogino K et al. J Biol Chem 2001 Aug 17;276(33):31376-87
PMID:11606526 - Characterization of Schizosaccharomyces pombe mcm7(+) and cdc23(+) (MCM10) and interactions with replication checkpoints.
Liang DT et al. Genetics 2001 Oct;159(2):471-86
PMID:10411894 - Cell cycle regulation of Dfp1, an activator of the Hsk1 protein kinase.
Brown GW et al. Proc Natl Acad Sci U S A 1999 Jul 20;96(15):8443-8
PMID:34608864 - DDK/Hsk1 phosphorylates and targets fission yeast histone deacetylase Hst4 for degradation to stabilize stalled DNA replication forks.
Aricthota S et al. Elife 2021 Oct 05;10
PMID:18505873 - Schizosaccharomyces pombe histone acetyltransferase Mst1 (KAT5) is an essential protein required for damage response and chromosome segregation.
Gómez EB et al. Genetics 2008 Jun;179(2):757-71
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:19422421 - Interactions between Swi1-Swi3, Mrc1 and S phase kinase, Hsk1 may regulate cellular responses to stalled replication forks in fission yeast.
Shimmoto M et al. Genes Cells 2009 Jun;14(6):669-82
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: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:19547744 - Evolution of phosphoregulation: comparison of phosphorylation patterns across yeast species.
Beltrao P et al. PLoS Biol 2009 Jun 16;7(6):e1000134
PMID:16263721 - Hsk1-Dfp1/Him1, the Cdc7-Dbf4 kinase in Schizosaccharomyces pombe, associates with Swi1, a component of the replication fork protection complex.
Matsumoto S et al. J Biol Chem 2005 Dec 30;280(52):42536-42
PMID:19443688 - Diverse roles of HP1 proteins in heterochromatin assembly and functions in fission yeast.
Fischer T et al. Proc Natl Acad Sci U S A 2009 Jun 02;106(22):8998-9003
PMID:22024164 - Multiple pathways can bypass the essential role of fission yeast Hsk1 kinase in DNA replication initiation.
Matsumoto S et al. J Cell Biol 2011 Oct 31;195(3):387-401
PMID:15643072 - Mcl1p is a polymerase alpha replication accessory factor important for S-phase DNA damage survival.
Williams DR et al. Eukaryot Cell 2005 Jan;4(1):166-77
PMID:36408920 - UniProt: the Universal Protein Knowledgebase in 2023.
UniProt Consortium Nucleic Acids Res 2023 Jan 06;51(D1):D523-D531
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:27320927 - Rif1 Regulates the Fate of DNA Entanglements during Mitosis.
Zaaijer S et al. Cell Rep 2016 Jun 28;16(1):148-160
PMID:12455694 - mcl1+, the Schizosaccharomyces pombe homologue of CTF4, is important for chromosome replication, cohesion, and segregation.
Williams DR et al. Eukaryot Cell 2002 Oct;1(5):758-73
PMID:10886372 - The Schizosaccharomyces pombe spo6+ gene encoding a nuclear protein with sequence similarity to budding yeast Dbf4 is required for meiotic second division and sporulation.
Nakamura T et al. Genes Cells 2000 Jun;5(6):463-79
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:18799612 - The Hsk1(Cdc7) replication kinase regulates origin efficiency.
Patel PK et al. Mol Biol Cell 2008 Dec;19(12):5550-8
PMID:23322785 - The fission yeast minichromosome maintenance (MCM)-binding protein (MCM-BP), Mcb1, regulates MCM function during prereplicative complex formation in DNA replication.
Santosa V et al. J Biol Chem 2013 Mar 08;288(10):6864-80
PMID:24656819 - Protein phosphatase 1 recruitment by Rif1 regulates DNA replication origin firing by counteracting DDK activity.
Davé A et al. Cell Rep 2014 Apr 10;7(1):53-61
PMID:27473316 - Characterization of a Novel MMS-Sensitive Allele of Schizosaccharomyces pombe mcm4.
Ranatunga NS et al. G3 (Bethesda) 2016 Oct 13;6(10):3049-3063
PMID:18180284 - Minichromosome maintenance proteins interact with checkpoint and recombination proteins to promote s-phase genome stability.
Bailis JM et al. Mol Cell Biol 2008 Mar;28(5):1724-38
PMID:21095590 - DDK phosphorylates checkpoint clamp component Rad9 and promotes its release from damaged chromatin.
Furuya K et al. Mol Cell 2010 Nov 24;40(4):606-18
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: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:31719112 - Checkpoint Regulation of Nuclear Tos4 Defines S Phase Arrest in Fission Yeast.
Kim SM et al. G3 (Bethesda) 2020 Jan 07;10(1):255-266
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:12161753 - The transcriptional program of meiosis and sporulation in fission yeast.
Mata J et al. Nat Genet 2002 Sep;32(1):143-7
PMID:10409743 - A fission yeast gene, him1(+)/dfp1(+), encoding a regulatory subunit for Hsk1 kinase, plays essential roles in S-phase initiation as well as in S-phase checkpoint control and recovery from DNA damage.
Takeda T et al. Mol Cell Biol 1999 Aug;19(8):5535-47
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: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:28069740 - Checkpoint-Independent Regulation of Origin Firing by Mrc1 through Interaction with Hsk1 Kinase.
Matsumoto S et al. Mol Cell Biol 2017 Apr 01;37(7)
PMID:18257517 - Phosphoproteome analysis of fission yeast.
Wilson-Grady JT et al. J Proteome Res 2008 Mar;7(3):1088-97
PMID:16199868 - Schizosaccharomyces pombe mst2+ encodes a MYST family histone acetyltransferase that negatively regulates telomere silencing.
Gómez EB et al. Mol Cell Biol 2005 Oct;25(20):8887-903
PMID:21518960 - Mrc1 marks early-firing origins and coordinates timing and efficiency of initiation in fission yeast.
Hayano M et al. Mol Cell Biol 2011 Jun;31(12):2380-91
PMID:15338237 - Schizosaccharomyces pombe replication protein Cdc45/Sna41 requires Hsk1/Cdc7 and Rad4/Cut5 for chromatin binding.
Dolan WP et al. Chromosoma 2004 Sep;113(3):145-56
PMID:26201080 - The DNA-Binding Domain of S. pombe Mrc1 (Claspin) Acts to Enhance Stalling at Replication Barriers.
Zech J et al. PLoS One 2015;10(7):e0132595
PMID:23862021 - The C-terminus of S. pombe DDK subunit Dfp1 is required for meiosis-specific transcription and cohesin cleavage.
Le AH et al. Biol Open 2013 Jul 15;2(7):728-38
PMID:27298342 - Identification of S-phase DNA damage-response targets in fission yeast reveals conservation of damage-response networks.
Willis NA et al. Proc Natl Acad Sci U S A 2016 Jun 28;113(26):E3676-85
PMID:20176980 - Fission yeast Hsk1 (Cdc7) kinase is required after replication initiation for induced mutagenesis and proper response to DNA alkylation damage.
Dolan WP et al. Genetics 2010 May;185(1):39-53
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:12604790 - The Cdc23 (Mcm10) protein is required for the phosphorylation of minichromosome maintenance complex by the Dfp1-Hsk1 kinase.
Lee JK et al. Proc Natl Acad Sci U S A 2003 Mar 04;100(5):2334-9
PMID:33313903 - Ribosome profiling reveals ribosome stalling on tryptophan codons and ribosome queuing upon oxidative stress in fission yeast.
Rubio A et al. Nucleic Acids Res 2021 Jan 11;49(1):383-399
PMID:7621824 - hsk1+, a Schizosaccharomyces pombe gene related to Saccharomyces cerevisiae CDC7, is required for chromosomal replication.
Masai H et al. EMBO J 1995 Jul 03;14(13):3094-104
PMID:16141239 - Endocytosis in fission yeast is spatially associated with the actin cytoskeleton during polarised cell growth and cytokinesis.
Gachet Y et al. J Cell Sci 2005 Sep 15;118(Pt 18):4231-42
PMID:14625560 - Hsk1-Dfp1 is required for heterochromatin-mediated cohesion at centromeres.
Bailis JM et al. Nat Cell Biol 2003 Dec;5(12):1111-6
PMID:12052858 - A conserved domain of Schizosaccharomyces pombe dfp1(+) is uniquely required for chromosome stability following alkylation damage during S phase.
Fung AD et al. Mol Cell Biol 2002 Jul;22(13):4477-90
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:21511999 - Comparative functional genomics of the fission yeasts.
Rhind N et al. Science 2011 May 20;332(6032):930-6