Gene Symbol: tko
Description: technical knockout
Alias: CG7925, Dmel\CG7925, EG:BACH59J11.1, MRP-S12, MRPS12, S12, Tko, l(1)3Ab, l(1)tko, mRpS12, mt-rps12, rps12, technical knockout, CG7925-PB, CG7925-PC, mitochondrial ribosomal protein S12, mitoribosomal protein S12, tko-PB, tko-PC
Species: fruit fly

Top Publications

  1. Kemppainen E, Fernández Ayala D, Galbraith L, O Dell K, Jacobs H. Phenotypic suppression of the Drosophila mitochondrial disease-like mutant tko(25t) by duplication of the mutant gene in its natural chromosomal context. Mitochondrion. 2009;9:353-63 pubmed publisher
    A mutation in the Drosophila gene technical knockout (tko(25t)), encoding mitoribosomal protein S12, phenocopies human mitochondrial disease...
  2. Lee J, Wu C. Electroconvulsive seizure behavior in Drosophila: analysis of the physiological repertoire underlying a stereotyped action pattern in bang-sensitive mutants. J Neurosci. 2002;22:11065-79 pubmed
  3. Shah Z, O Dell K, Miller S, An X, Jacobs H. Metazoan nuclear genes for mitoribosomal protein S12. Gene. 1997;204:55-62 pubmed
    We have characterized nuclear genes for mitoribosomal protein S12 (mt-rps12) a major component of the ribosomal accuracy centre, in human, mouse and Drosophila melanogaster...
  4. Royden C, Pirrotta V, Jan L. The tko locus, site of a behavioral mutation in D. melanogaster, codes for a protein homologous to prokaryotic ribosomal protein S12. Cell. 1987;51:165-73 pubmed
    The tko (technical knockout) mutation is one of a family of behavioral mutations that cause "bang sensitivity" in D. melanogaster. Using P-element-mediated transformation, we show that a 3...
  5. Fergestad T, Bostwick B, Ganetzky B. Metabolic disruption in Drosophila bang-sensitive seizure mutants. Genetics. 2006;173:1357-64 pubmed
    ..Our data implicate cellular metabolism in regulating seizure susceptibility and suggest that differential sensitivity of neuronal subtypes to metabolic changes underlies distinct types of seizure activity. ..
  6. Pavlidis P, Tanouye M. Seizures and failures in the giant fiber pathway of Drosophila bang-sensitive paralytic mutants. J Neurosci. 1995;15:5810-9 pubmed
    ..This hypothesis is supported by the finding that, in double mutant combination with mlenapts, which suppresses behavioral bang sensitivity, DLM failures, but not seizures, were reduced. ..
  7. Chen S, Oliveira M, Sanz A, Kemppainen E, Fukuoh A, Schlicht B, et al. A cytoplasmic suppressor of a nuclear mutation affecting mitochondrial functions in Drosophila. Genetics. 2012;192:483-93 pubmed publisher
    ..cytoplasmic suppression of the mitochondrial disease-like phenotype of the Drosophila melanogaster nuclear mutant tko(25t), which includes developmental delay, seizure sensitivity, and defective male courtship...
  8. Judd B, Shen M, Kaufman T. The anatomy and function of a segment of the X chromosome of Drosophila melanogaster. Genetics. 1972;71:139-56 pubmed
    ..The data clearly show one functional group per chromomere. It is postulated that a chromomere is one cistron within which much of the DNA is regulatory in function. ..
  9. Fergestad T, Olson L, Patel K, Miller R, Palladino M, Ganetzky B. Neuropathology in Drosophila mutants with increased seizure susceptibility. Genetics. 2008;178:947-56 pubmed publisher
    ..These results emphasize the importance of proper cellular metabolism in maintaining both the activity and viability of neurons. ..

More Information


  1. Toivonen J, O Dell K, Petit N, Irvine S, Knight G, Lehtonen M, et al. Technical knockout, a Drosophila model of mitochondrial deafness. Genetics. 2001;159:241-54 pubmed
    ..To develop animal models of such disorders, we have manipulated the nuclear gene for mitochondrial ribosomal protein S12 in Drosophila (technical knockout, tko)...
  2. Engel J, Wu C. Altered mechanoreceptor response in Drosophila bang-sensitive mutants. J Comp Physiol A. 1994;175:267-78 pubmed
    ..DNA sequence analysis of tko and eas has indicated that they encode apparently unrelated biochemical products...
  3. Shannon M, Kaufman T, Shen M, Judd B. Lethality patterns and morphology of selected lethal and semi-lethal mutations in the zeste-white region of Drosophila melanogaster. Genetics. 1972;72:615-38 pubmed
    ..These results provide additional evidence that only one function is associated with each chromomere.-The results of the lethality pattern analysis are also compared with previous studies of lethal mutants of Drosophila. ..
  4. Song J, Tanouye M. Seizure suppression by shakB2, a gap junction mutation in Drosophila. J Neurophysiol. 2006;95:627-35 pubmed
    ..Seizures caused by easily shocked (eas) and technical knockout (tko) mutations are partially suppressed by shakB2...
  5. Goldberg M, Colvin R, Mellin A. The Drosophila zeste locus is nonessential. Genetics. 1989;123:145-55 pubmed
    ..This suggests that the zeste protein normally is either a very weak transcription factor, or that its function can be substituted by that of other regulatory proteins. ..
  6. Nomura T, Kurokawa N. Comparative study on germ cell mutation induced by urethane (ethyl carbamate) gas and X-rays in Drosophila melanogaster. Jpn J Cancer Res. 1997;88:461-7 pubmed
    ..Urethane-induced mutations were strikingly non-random with two hot spots at zw-1 and zw-2, whereas the distribution of X-ray-induced mutations was more nearly random. ..
  7. Burg M, Wu C. Mechanical and temperature stressor-induced seizure-and-paralysis behaviors in Drosophila bang-sensitive mutants. J Neurogenet. 2012;26:189-97 pubmed publisher
    ..The authors found that each of the bang-sensitive mutants described in this paper (bas, bss, eas, and tko) also displayed similar behavioral repertoires upon exposure to either high or low temperature...
  8. Tower J. Mitochondrial maintenance failure in aging and role of sexual dimorphism. Arch Biochem Biophys. 2015;576:17-31 pubmed publisher
    ..The data suggest that metazoan life span interventions may act through a common hormesis mechanism involving liver UPRmt, mitochondrial maintenance and sexual differentiation. ..
  9. Paradi E, Vogel E, Szilagyi E. Effect of storage and dose on MMS-induced deletions. Complementation analysis of X-chromosomal recessive lethals in the zeste-white and maroon-like regions of Drosophila melanogaster. Mutat Res. 1983;111:145-59 pubmed
    ..An interesting aspect for further analysis is the apparent infrequency in the zeste-white region of alkylation-induced chromosomal breakage, as observed by various investigators for MMS, EMS and MNNG. ..
  10. Stone B, Evans L, Coleman J, Kuebler D. Genetic and pharmacological manipulations that alter metabolism suppress seizure-like activity in Drosophila. Brain Res. 2013;1496:94-103 pubmed publisher
    ..the atsugari (atu) mutation into three of the BS mutants, easily shocked (eas), bang senseless (bss), and technical knockout (tko)...
  11. Crandall K, Templeton A. Empirical tests of some predictions from coalescent theory with applications to intraspecific phylogeny reconstruction. Genetics. 1993;134:959-69 pubmed
  12. Clark A. Inference of haplotypes from PCR-amplified samples of diploid populations. Mol Biol Evol. 1990;7:111-22 pubmed
    ..The algorithm also applies to the problem of inferring haplotype frequencies of closely linked restriction-site polymorphisms. ..
  13. Toivonen J, Walker G, Martinez Diaz P, Bjedov I, Driege Y, Jacobs H, et al. No influence of Indy on lifespan in Drosophila after correction for genetic and cytoplasmic background effects. PLoS Genet. 2007;3:e95 pubmed
    ..In addition, we saw no effects on lifespan of expression knockdown of the Indy orthologues nac-2 and nac-3 in the nematode Caenorhabditis elegans. ..
  14. Liu C, Lim J. Complementation analysis of methyl methane-sulfonate-induced recessive lethal mutations in the zeste-white region of the X chromosome of Drosophila melanogaster. Genetics. 1975;79:601-11 pubmed
    ..One of these single-site mutations was in qt, 2 in tko, 18 in zw-1, 12 in zw-8, 6 in zw-4, 3 in zw-10, 3 in zw-13, 21 in zw-2, 7 in zw-3, 5 in zw-6, 6 in zw-12, 1 in zw-7,..
  15. Amikura R, Kashikawa M, Nakamura A, Kobayashi S. Presence of mitochondria-type ribosomes outside mitochondria in germ plasm of Drosophila embryos. Proc Natl Acad Sci U S A. 2001;98:9133-8 pubmed
    ..reveals that these ribosomes include both mitochondrial rRNAs and at least two mitochondrial ribosomal proteins (S12 and L7/L12)...
  16. Hardin P. From biological clock to biological rhythms. Genome Biol. 2000;1:REVIEWS1023 pubmed
    ..Completion of whole genome sequences and the use of differential gene expression technology will hasten the discovery of the clock output pathways that control diverse rhythmic phenomena. ..
  17. Kemppainen K, Kemppainen E, Jacobs H. The alternative oxidase AOX does not rescue the phenotype of tko25t mutant flies. G3 (Bethesda). 2014;4:2013-21 pubmed publisher
    A point mutation [technical knockout(25t) (tko(25t))] in the Drosophila gene coding for mitoribosomal protein S12 generates a phenotype of developmental delay and bang sensitivity...
  18. Begun D, Aquadro C. Levels of naturally occurring DNA polymorphism correlate with recombination rates in D. melanogaster. Nature. 1992;356:519-20 pubmed
    ..Hitch-hiking thus seems to occur over a large fraction of the Drosophila genome and may constitute a major constraint on levels of genetic variation in nature. ..
  19. Tan J, Lin F, Tanouye M. Potassium bromide, an anticonvulsant, is effective at alleviating seizures in the Drosophila bang-sensitive mutant bang senseless. Brain Res. 2004;1020:45-52 pubmed
    ..This methodology suggests that the Drosophila system can potentially be a powerful instrument for assaying and testing new compounds with anticonvulsant properties. ..
  20. Whelan J, Burke B, Rice A, Tong M, Kuebler D. Sensitivity to seizure-like activity in Drosophila following acute hypoxia and hypercapnia. Brain Res. 2010;1316:120-8 pubmed publisher
    ..The increased sensitivity to both hypoxia and hypercapnia in these BS mutants suggests possible physiological defects that may underlie seizure susceptibility. ..
  21. Sen A, Damm V, Cox R. Drosophila clueless is highly expressed in larval neuroblasts, affects mitochondrial localization and suppresses mitochondrial oxidative damage. PLoS ONE. 2013;8:e54283 pubmed publisher
    ..Mutations in two other genes encoding mitochondrial proteins, technical knockout and stress sensitive B, do not cause neuroblast mitochondrial mislocalization, even though technical ..
  22. Wei Y, Mondal S, Mouawad R, Wilczyński B, Henry R, Arnosti D. Genome-Wide Analysis of Drosophila RBf2 Protein Highlights the Diversity of RB Family Targets and Possible Role in Regulation of Ribosome Biosynthesis. G3 (Bethesda). 2015;5:1503-15 pubmed publisher
    ..The presence of unique RBf2 targets suggest that evolutionary appearance of this corepressor represents the acquisition of potentially novel roles in gene regulation for the RB family. ..
  23. Mandavilli B, Santos J, Van Houten B. Mitochondrial DNA repair and aging. Mutat Res. 2002;509:127-51 pubmed
    ..Mitochondrial DNA damage, if not repaired, leads to disruption of electron transport chain and production of more ROS. This vicious cycle of ROS production and mtDNA damage ultimately leads to energy depletion in the cell and apoptosis. ..
  24. Jacobs H, Fernandez Ayala D, Manjiry S, Kemppainen E, Toivonen J, O Dell K. Mitochondrial disease in flies. Biochim Biophys Acta. 2004;1659:190-6 pubmed
    The Drosophila mutant technical knockout (tko), affecting the mitochondrial protein synthetic apparatus, exhibits respiratory chain deficiency and a phenotype resembling various features of mitochondrial disease in humans (paralytic ..
  25. Emmons S, Lipton J. Genetic basis of male sexual behavior. J Neurobiol. 2003;54:93-110 pubmed
    ..It can also help us to gain an appreciation of how behavior is encoded by genes and their regulatory sequences. ..
  26. Ganetzky B, Wu C. Indirect Suppression Involving Behavioral Mutants with Altered Nerve Excitability in DROSOPHILA MELANOGASTER. Genetics. 1982;100:597-614 pubmed
    ..Genetic interactions of this type help reveal functional relationships between different behavioral mutants and suggest ways of isolating new mutants with altered excitable membranes. ..
  27. Georgiev P, Okkenhaug H, Drews A, Wright D, Lambert S, Flick M, et al. TRPM channels mediate zinc homeostasis and cellular growth during Drosophila larval development. Cell Metab. 2010;12:386-397 pubmed publisher
    ..We propose that regulation of Zn(2+) homeostasis through dTRPM channels is required to support molecular processes that mediate class I PI3K-regulated cell growth. ..
  28. Lim J, Snyder L. Cytogenetic and complementation analyses of recessive lethal mutations induced in the X chromosome of Drosophila by three alkylating agents. Genet Res. 1974;24:1-10 pubmed
  29. Montgomery E, Huang S, Langley C, Judd B. Chromosome rearrangement by ectopic recombination in Drosophila melanogaster: genome structure and evolution. Genetics. 1991;129:1085-98 pubmed
  30. Datta S, Kankel D. l(1)trol and l(1)devl, loci affecting the development of the adult central nervous system in Drosophila melanogaster. Genetics. 1992;130:523-37 pubmed
    ..Two of these, including one that is allelic to l(1)zw1, were chosen for further analysis. ..
  31. Chihara T, Luginbuhl D, Luo L. Cytoplasmic and mitochondrial protein translation in axonal and dendritic terminal arborization. Nat Neurosci. 2007;10:828-37 pubmed
    ..Our study highlights different demands of protein translation for the development and maintenance of axons and dendrites. ..
  32. Garcia Bellido A, Robbins L. Viability of Female Germ-Line Cells Homozygous for Zygotic Lethals in DROSOPHILA MELANOGASTER. Genetics. 1983;103:235-47 pubmed
    ..The abundance of genes whose products are required for oogenesis, whose products are required in the oocyte, and whose activity is required during zygotic development is discussed. ..
  33. Thierry Mieg D. Paralog, a control mutant in Drosophila melanogaster. Genetics. 1982;100:209-37 pubmed
    ..We interpret the data by postulating that the expression of sets of dispersed genes might be controlled by the local topology of the chromosome, itself constrained by pairing of dispersed repeated elements. We call the mutation paralog. ..
  34. Lin H, Wolfner M. Cloning and analysis of fs(1) Ya, a maternal effect gene required for the initiation of Drosophila embryogenesis. Mol Gen Genet. 1989;215:257-65 pubmed
    ..The transcript is not detected in later stages of embryonic development. This expression pattern correlates closely with the genetic and developmental characteristics expected of the fs(1) Ya gene product. ..
  35. Lynch M, Crease T. The analysis of population survey data on DNA sequence variation. Mol Biol Evol. 1990;7:377-94 pubmed publisher
    ..Generally, the degree of population subdivision at the nucleotide level is comparable with that at the haplotype level, but significant differences do arise as a result of inequalities in the genetic distances between haplotypes...
  36. Lee J, Wu C. Genetic modifications of seizure susceptibility and expression by altered excitability in Drosophila Na(+) and K(+) channel mutants. J Neurophysiol. 2006;96:2465-78 pubmed
    ..Our results suggest that individual seizure repertoire components reflect different neural network activities that could be differentially altered by mutations of specific ion channel subunits. ..