Gene Symbol: tor2
Description: phosphatidylinositol kinase Tor2
Alias: SPBC646.01c, phosphatidylinositol kinase Tor2
Species: fission yeast

Top Publications

  1. Weisman R, Roitburg I, Schonbrun M, Harari R, Kupiec M. Opposite effects of tor1 and tor2 on nitrogen starvation responses in fission yeast. Genetics. 2007;175:1153-62 pubmed
    ..In the fission yeast two TOR homologs are present. tor1(+) is required for starvation and stress responses, while tor2(+) is essential...
  2. Urano J, Sato T, Matsuo T, Otsubo Y, Yamamoto M, Tamanoi F. Point mutations in TOR confer Rheb-independent growth in fission yeast and nutrient-independent mammalian TOR signaling in mammalian cells. Proc Natl Acad Sci U S A. 2007;104:3514-9 pubmed
    ..In this article, we report identification of 22 single amino acid changes in the Tor2 protein that enable growth in the absence of Rhb1p. These mutants also exhibit decreased mating efficiency...
  3. Hayashi T, Hatanaka M, Nagao K, Nakaseko Y, Kanoh J, Kokubu A, et al. Rapamycin sensitivity of the Schizosaccharomyces pombe tor2 mutant and organization of two highly phosphorylated TOR complexes by specific and common subunits. Genes Cells. 2007;12:1357-70 pubmed
    ..of Schizosaccharomyces pombe temperature-sensitive strains led to the isolation of a nutrient-insensitive mutant, tor2-287...
  4. Nakashima A, Sato T, Tamanoi F. Fission yeast TORC1 regulates phosphorylation of ribosomal S6 proteins in response to nutrients and its activity is inhibited by rapamycin. J Cell Sci. 2010;123:777-86 pubmed publisher
    ..In Schizosaccharomyces pombe, TORC1, which contains Tor2, plays crucial roles in nutrient response. Here we find a nitrogen-regulated phosphoprotein, p27, in S...
  5. Uritani M, Hidaka H, Hotta Y, Ueno M, Ushimaru T, Toda T. Fission yeast Tor2 links nitrogen signals to cell proliferation and acts downstream of the Rheb GTPase. Genes Cells. 2006;11:1367-79 pubmed
    ..Yeast contains two related proteins, Tor1 and Tor2. In fission yeast, Tor1 is dispensable for normal growth but is involved in amino acid uptake and cell survival ..
  6. Ikai N, Nakazawa N, Hayashi T, Yanagida M. The reverse, but coordinated, roles of Tor2 (TORC1) and Tor1 (TORC2) kinases for growth, cell cycle and separase-mediated mitosis in Schizosaccharomyces pombe. Open Biol. 2011;1:110007 pubmed publisher
    ..These results were obtained using the previously isolated TORC1 mutant tor2-L2048S in the phosphatidyl inositol kinase (PIK) domain and a new TORC2 mutant tor1-L2045D, which harbours a ..
  7. Valbuena N, Moreno S. TOR and PKA pathways synergize at the level of the Ste11 transcription factor to prevent mating and meiosis in fission yeast. PLoS ONE. 2010;5:e11514 pubmed publisher
    ..Inhibition of Tor2 signaling results in the induction of genes involved in sexual differentiation, and the cells undergo mating and ..
  8. Alvarez B, Moreno S. Fission yeast Tor2 promotes cell growth and represses cell differentiation. J Cell Sci. 2006;119:4475-85 pubmed
    ..The molecular mechanisms underlying this response are not fully understood. Here, we demonstrate that Tor2, a fission yeast member of the TOR protein kinase family, is central to controlling the switch between cell growth ..
  9. Nakashima A, Otsubo Y, Yamashita A, Sato T, Yamamoto M, Tamanoi F. Psk1, an AGC kinase family member in fission yeast, is directly phosphorylated and controlled by TORC1 and functions as S6 kinase. J Cell Sci. 2012;125:5840-9 pubmed publisher
    ..In the fission yeast Schizosaccharomyces pombe, TOR complex 1 (TORC1), which includes Tor2 as a catalytic subunit, manages the switch between cell proliferation and differentiation by sensing nutrient ..

More Information


  1. Takahara T, Maeda T. TORC1 of fission yeast is rapamycin-sensitive. Genes Cells. 2012;17:698-708 pubmed publisher
    ..These results indicate that rapamycin does inhibit TORC1 in S. pombe, but the inhibition is not sufficient to cause a growth defect. These findings establish a universal action of rapamycin on TORC1 inhibition. ..
  2. Ikeda K, Morigasaki S, Tatebe H, Tamanoi F, Shiozaki K. Fission yeast TOR complex 2 activates the AGC-family Gad8 kinase essential for stress resistance and cell cycle control. Cell Cycle. 2008;7:358-64 pubmed
    ..These results strongly suggest that the TORC2-Gad8 pathway has multiple physiological functions in cellular stress resistance and cell cycle progression at both G(1)/S and G(2)/M transitions. ..
  3. Matsuo T, Otsubo Y, Urano J, Tamanoi F, Yamamoto M. Loss of the TOR kinase Tor2 mimics nitrogen starvation and activates the sexual development pathway in fission yeast. Mol Cell Biol. 2007;27:3154-64 pubmed
    Fission yeast has two TOR (target of rapamycin) kinases, namely Tor1 and Tor2. Tor1 is required for survival under stressed conditions, proper G(1) arrest, and sexual development. In contrast, Tor2 is essential for growth...
  4. Ma N, Ma Y, Nakashima A, Kikkawa U, Furuyashiki T. The Loss of Lam2 and Npr2-Npr3 Diminishes the Vacuolar Localization of Gtr1-Gtr2 and Disinhibits TORC1 Activity in Fission Yeast. PLoS ONE. 2016;11:e0156239 pubmed publisher
    ..These findings suggest that Lam2 and Npr2-Npr3 function together as a tether for GDP-bound Gtr1 to the vacuolar membrane, thereby suppressing TORC1 activity for multiple cellular functions. ..
  5. Otsubo Y, Yamashita A, Ohno H, Yamamoto M. S. pombe TORC1 activates the ubiquitin-proteasomal degradation of the meiotic regulator Mei2 in cooperation with Pat1 kinase. J Cell Sci. 2014;127:2639-46 pubmed publisher
    ..Based on these data, we propose that TORC1 contributes to the suppression of sexual development by phosphorylating Mei2, in addition to controlling the cellular metabolic status. ..
  6. Otsubo Y, Nakashima A, Yamamoto M, Yamashita A. TORC1-Dependent Phosphorylation Targets in Fission Yeast. Biomolecules. 2017;7: pubmed publisher
    ..TORC1, which contains Tor2 as a catalytic subunit, promotes vegetative growth and represses sexual differentiation in nutrient-rich conditions,..
  7. Cohen A, Kupiec M, Weisman R. Gad8 Protein Is Found in the Nucleus Where It Interacts with the MluI Cell Cycle Box-binding Factor (MBF) Transcriptional Complex to Regulate the Response to DNA Replication Stress. J Biol Chem. 2016;291:9371-81 pubmed publisher
    ..Taken together, our findings highlight critical functions of TORC2 in the nucleus and suggest a role in surviving DNA replication stress via transcriptional regulation of MBF target genes. ..
  8. Saitoh S, Mori A, Uehara L, Masuda F, Soejima S, Yanagida M. Mechanisms of expression and translocation of major fission yeast glucose transporters regulated by CaMKK/phosphatases, nuclear shuttling, and TOR. Mol Biol Cell. 2015;26:373-86 pubmed publisher
    ..These results clarify the evolutionarily conserved molecular mechanisms underlying glucose homeostasis, which are essential for preventing hyperglycemia in humans. ..
  9. Ma N, Liu Q, Zhang L, Henske E, Ma Y. TORC1 signaling is governed by two negative regulators in fission yeast. Genetics. 2013;195:457-68 pubmed publisher
    ..Furthermore, we showed that Tor2 controls amino acid homeostasis at the transcriptional and post-transcriptional levels...
  10. Laor D, Cohen A, Pasmanik Chor M, Oron Karni V, Kupiec M, Weisman R. Isp7 is a novel regulator of amino acid uptake in the TOR signaling pathway. Mol Cell Biol. 2014;34:794-806 pubmed publisher
    ..Taken together, our findings suggest a central role for Isp7 in amino acid homeostasis and the presence of isp7(+)-dependent regulatory loops that affect both TORC1 and TORC2. ..
  11. Valbuena N, Rozalén A, Moreno S. Fission yeast TORC1 prevents eIF2? phosphorylation in response to nitrogen and amino acids via Gcn2 kinase. J Cell Sci. 2012;125:5955-9 pubmed publisher
    ..In this study, we show that Tor2, as part of the TORC1 complex, prevents the phosphorylation of eIF2? in cells growing in the presence of nitrogen ..
  12. Ma Y, Ma N, Liu Q, Qi Y, Manabe R, Furuyashiki T. Tor Signaling Regulates Transcription of Amino Acid Permeases through a GATA Transcription Factor Gaf1 in Fission Yeast. PLoS ONE. 2015;10:e0144677 pubmed publisher
    In the fission yeast, two Tor isoforms, Tor1 and Tor2, oppositely regulate gene expression of amino acid permeases...
  13. Valbuena N, Guan K, Moreno S. The Vam6 and Gtr1-Gtr2 pathway activates TORC1 in response to amino acids in fission yeast. J Cell Sci. 2012;125:1920-8 pubmed publisher
    ..Our data demonstrate the existence of an evolutionarily conserved pathway with the Vam6 and Gtr1-Gtr2 pathway activating TORC1, which in turns stimulates cell growth and inhibits sexual differentiation. ..
  14. Urano J, Comiso M, Guo L, Aspuria P, Deniskin R, Tabancay A, et al. Identification of novel single amino acid changes that result in hyperactivation of the unique GTPase, Rheb, in fission yeast. Mol Microbiol. 2005;58:1074-86 pubmed
    ..By using the hyperactive mutant, Rhb1(K120R), we have been able to demonstrate that Rhb1 interacts with Tor2, one of the two S. pombe TOR (Target of Rapamycin) proteins...
  15. Murai T, Nakase Y, Fukuda K, Chikashige Y, Tsutsumi C, Hiraoka Y, et al. Distinctive responses to nitrogen starvation in the dominant active mutants of the fission yeast Rheb GTPase. Genetics. 2009;183:517-27 pubmed publisher
    ..We postulate that the signaling cascade may branch below Rhb1 or Tor2 and regulate the amino acid uptake and response to nitrogen starvation independently.
  16. Nakase Y, Nakase M, Kashiwazaki J, Murai T, Otsubo Y, Mabuchi I, et al. The fission yeast ?-arrestin-like protein Any1 is involved in TSC-Rheb signaling and the regulation of amino acid transporters. J Cell Sci. 2013;126:3972-81 pubmed publisher
    ..Physical interaction of Pub1 and Any1 is more stable in ?tsc2 cells than in wild-type cells and is independent of Tor2 activity...
  17. Liu Q, Ma Y, Zhou X, Furuyashiki T. Constitutive Tor2 Activity Promotes Retention of the Amino Acid Transporter Agp3 at Trans-Golgi/Endosomes in Fission Yeast. PLoS ONE. 2015;10:e0139045 pubmed publisher
    ..There are two TOR isoforms in fission yeast, Tor1 and Tor2. Whereas tor1 deletion did not affect the Torin-1-induced transfer of Agp3-YFP, Tor2 inhibition using a temperature-..
  18. Chica N, Rozalén A, Pérez Hidalgo L, Rubio A, Novak B, Moreno S. Nutritional Control of Cell Size by the Greatwall-Endosulfine-PP2A·B55 Pathway. Curr Biol. 2016;26:319-30 pubmed publisher
    ..Given the conservation of this pathway, it is reasonable to assume that this mechanism operates in higher eukaryotes, as well. ..
  19. Pataki E, Weisman R, Sipiczki M, Miklos I. fhl1 gene of the fission yeast regulates transcription of meiotic genes and nitrogen starvation response, downstream of the TORC1 pathway. Curr Genet. 2017;63:91-101 pubmed publisher
    ..the majority of these genes belonged to the nitrogen starvation response, the possible cooperation of fhl1 and tor2 was also investigated...