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Daniel Portnoy

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Paolo S Manzanillo; Daniel A Portnoy; Michael Shiloh; Jeffery S Cox
Mycobacterium tuberculosis activates the DNA-dependent cytosolic surveillance pathway within macrophages.
Daniel A Portnoy
Yogi Berra, Forrest Gump, and the discovery of Listeria actin comet tails.
Jody A Melton-Witt; Daniel A Portnoy; Susanne M Rafelski; Anna I Bakardjiev
Oral infection with signature-tagged Listeria monocytogenes reveals organ-specific growth and dissemination routes in guinea pigs.

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All Publications

2012: Paolo S Manzanillo; Daniel A Portnoy; Michael Shiloh; Jeffery S Cox
Mycobacterium tuberculosis activates the DNA-dependent cytosolic surveillance pathway within macrophages.
Cell host & microbe 2012;11(5):469-80.
2012: Daniel A Portnoy
Yogi Berra, Forrest Gump, and the discovery of Listeria actin comet tails.
Molecular biology of the cell 2012;23(7):1141-5.
2012: Jody A Melton-Witt; Daniel A Portnoy; Susanne M Rafelski; Anna I Bakardjiev
Oral infection with signature-tagged Listeria monocytogenes reveals organ-specific growth and dissemination routes in guinea pigs.
Infection and immunity 2012;80(2):720-32.
2012: Chelsea E Witte; Joshua J Woodward; Kristina A Archer; Chris S Rae; John-Demian Sauer; Daniel A Portnoy
Innate immune pathways triggered by Listeria monocytogenes and their role in the induction of cell-mediated immunity.
Advances in immunology 2012;113():135-56.
2011: Chris S Rae; Paul C Adamson; Aimee Geissler; Daniel A Portnoy
Mutations of the Listeria monocytogenes peptidoglycan N-deacetylase and O-acetylase result in enhanced lysozyme sensitivity, bacteriolysis, and hyperinduction of innate immune pathways.
Infection and immunity 2011;79(9):3596-606.
2011: John-Demian Sauer; Kristina A Archer; Thomas P Burke; Bill Hanson; Peter Lauer; Sabine Pereyre; Daniel A Portnoy
Listeria monocytogenes engineered to activate the Nlrc4 inflammasome are severely attenuated and are poor inducers of protective immunity.
Proceedings of the National Academy of Sciences of the United States of America 2011;108(30):12419-24.
2011: John-Demian Sauer; Katia Sotelo-Troha; Jakob von Moltke; Kathryn M Monroe; Chris S Rae; Sky W Brubaker; Mamoru Hyodo; Yoshihiro Hayakawa; Joshua J Woodward; Daniel A Portnoy; Russell E Vance
The N-ethyl-N-nitrosourea-induced Goldenticket mouse mutant reveals an essential function of Sting in the in vivo interferon response to Listeria monocytogenes and cyclic dinucleotides.
Infection and immunity 2011;79(2):688-94.
2010: Joshua J Woodward; Anthony T Iavarone; Daniel A Portnoy
c-di-AMP secreted by intracellular Listeria monocytogenes activates a host type I interferon response.
Science (New York, N.Y.) 2010;328(5986):1703-5.
2010: John-Demian Sauer; Chelsea E Witte; Jason Zemansky; Bill Hanson; Peter Lauer; Daniel A Portnoy
Listeria monocytogenes triggers AIM2-mediated pyroptosis upon infrequent bacteriolysis in the macrophage cytosol.
Cell host & microbe 2010;7(5):412-9.
2010: Nicole Meyer-Morse; Jennifer R Robbins; Chris S Rae; Sofia N Mochegova; Michele S Swanson; Zijiang Zhao; Herbert W Virgin; Daniel A Portnoy
Listeriolysin O is necessary and sufficient to induce autophagy during Listeria monocytogenes infection.
PloS one 2010;5(1):e8610.
2009: Keith S Bahjat; Nicole Meyer-Morse; Edward E Lemmens; Jessica A Shugart; Thomas W Dubensky; Dirk G Brockstedt; Daniel A Portnoy
Suppression of cell-mediated immunity following recognition of phagosome-confined bacteria.
PLoS pathogens 2009;5(9):e1000568.
2009: Russell E Vance; Ralph R Isberg; Daniel A Portnoy
Patterns of pathogenesis: discrimination of pathogenic and nonpathogenic microbes by the innate immune system.
Cell host & microbe 2009;6(1):10-21.
2009: Gregory T Crimmins; Michael W Schelle; Anat Herskovits; Peggy P Ni; Benjamin C Kline; Nicole Meyer-Morse; Anthony T Iavarone; Daniel A Portnoy
Listeria monocytogenes 6-Phosphogluconolactonase mutants induce increased activation of a host cytosolic surveillance pathway.
Infection and immunity 2009;77(7):3014-22.
2009: Jason Zemansky; Benjamin C Kline; Joshua J Woodward; Jess H Leber; Hélène Marquis; Daniel A Portnoy
Development of a mariner-based transposon and identification of Listeria monocytogenes determinants, including the peptidyl-prolyl isomerase PrsA2, that contribute to its hemolytic phenotype.
Journal of bacteriology 2009;191(12):3950-64.
2008: Gregory T Crimmins; Anat Herskovits; Kai Rehder; Kelsey E Sivick; Peter Lauer; Thomas W Dubensky; Daniel A Portnoy
Listeria monocytogenes multidrug resistance transporters activate a cytosolic surveillance pathway of innate immunity.
Proceedings of the National Academy of Sciences of the United States of America 2008;105(29):10191-6.
2008: Julie P M Viala; Sofia N Mochegova; Nicole Meyer-Morse; Daniel A Portnoy
A bacterial pore-forming toxin forms aggregates in cells that resemble those associated with neurodegenerative diseases.
Cellular microbiology 2008;10(4):985-93.
2008: Jess H Leber; Gregory T Crimmins; Sridharan Raghavan; Nicole P Meyer-Morse; Jeffery S Cox; Daniel A Portnoy
Distinct TLR- and NLR-mediated transcriptional responses to an intracellular pathogen.
PLoS pathogens 2008;4(1):e6.
2007: Keith Bahjat; Rodney A Prell; Heather E Allen; Weiqun Liu; Edward E Lemmens; Meredith L Leong; Daniel A Portnoy; Thomas W Dubensky; Dirk G Brockstedt; Martin A Giedlin
Activation of immature hepatic NK cells as immunotherapy for liver metastatic disease.
Journal of immunology (Baltimore, Md. : 1950) 2007;179(11):7376-84.
2007: Pamela Schnupf; Jianmin Zhou; Alexander Varshavsky; Daniel A Portnoy
Listeriolysin O secreted by Listeria monocytogenes into the host cell cytosol is degraded by the N-end rule pathway.
Infection and immunity 2007;75(11):5135-47.
2007: Pamela Schnupf; Daniel A Portnoy
Listeriolysin O: a phagosome-specific lysin.
Microbes and infection / Institut Pasteur 2007;9(10):1176-87.
2007: Anat Herskovits; Victoria Auerbuch; Daniel A Portnoy
Bacterial ligands generated in a phagosome are targets of the cytosolic innate immune system.
PLoS pathogens 2007;3(3):e51.
2006: Keith Bahjat; Weiqun Liu; Edward E Lemmens; Stephen P Schoenberger; Daniel A Portnoy; Thomas W Dubensky; Dirk G Brockstedt
Cytosolic entry controls CD8+-T-cell potency during bacterial infection.
Infection and immunity 2006;74(11):6387-97.
2006: Anna I Bakardjiev; Julie A Theriot; Daniel A Portnoy
Listeria monocytogenes traffics from maternal organs to the placenta and back.
PLoS pathogens 2006;2(6):e66.
2006: Pamela Schnupf; Daniel A Portnoy; Amy L Decatur
Phosphorylation, ubiquitination and degradation of listeriolysin O in mammalian cells: role of the PEST-like sequence.
Cellular microbiology 2006;8(2):353-64.
2006: Koichi Tabeta; Kasper Hoebe; edith janssen; Xin Du; Philippe Georgel; Karine Crozat; Suzanne Mudd; Navjiwan Mann; Sosathya Sovath; Jason Goode; Louis Shamel; Anat Herskovits; Daniel A Portnoy; Michael Cooke; Lisa Tarantino; Timothy Wiltshire; Benjamin E Steinberg; Sergio Grinstein; Bruce Beutler
The Unc93b1 mutation 3d disrupts exogenous antigen presentation and signaling via Toll-like receptors 3, 7 and 9.
Nature immunology 2006;7(2):156-64.
2005: Luisa W Cheng; Julie P M Viala; Nico Stuurman; Ursula Wiedemann; Ronald D Vale; Daniel A Portnoy
Use of RNA interference in Drosophila S2 cells to identify host pathways controlling compartmentalization of an intracellular pathogen.
Proceedings of the National Academy of Sciences of the United States of America 2005;102(38):13646-51.
2005: Dirk G Brockstedt; Keith Bahjat; Martin A Giedlin; Weiqun Liu; Meredith L Leong; William Luckett; Y Gao; Pamela Schnupf; D Kapadia; G Castro; J Y H Lim; A Sampson-Johannes; Anat Herskovits; Adonis Stassinopoulos; H G Archie Bouwer; J E Hearst; Daniel A Portnoy; David N Cook; Thomas W Dubensky
Killed but metabolically active microbes: a new vaccine paradigm for eliciting effector T-cell responses and protective immunity.
Nature medicine 2005;11(8):853-60.
2005: Anna I Bakardjiev; Brian A Stacy; Daniel A Portnoy
Growth of Listeria monocytogenes in the guinea pig placenta and role of cell-to-cell spread in fetal infection.
The Journal of infectious diseases 2005;191(11):1889-97.
2005: Daniel A Portnoy
Manipulation of innate immunity by bacterial pathogens.
Current opinion in immunology 2005;17(1):25-8.
2004: Dirk G Brockstedt; Martin A Giedlin; Meredith L Leong; Keith Bahjat; Yi Gao; William Luckett; Weiqun Liu; David N Cook; Daniel A Portnoy; Thomas W Dubensky
Listeria-based cancer vaccines that segregate immunogenicity from toxicity.
Proceedings of the National Academy of Sciences of the United States of America 2004;101(38):13832-7.
2004: Victoria Auerbuch; Dirk G Brockstedt; Nicole Meyer-Morse; Mary O'Riordan; Daniel A Portnoy
Mice lacking the type I interferon receptor are resistant to Listeria monocytogenes.
The Journal of experimental medicine 2004;200(4):527-33.
2004: Ramona L McCaffrey; Paul Fawcett; Mary O'Riordan; Kyung-Dall Lee; Edward A Havell; Patrick O Brown; Daniel A Portnoy
A specific gene expression program triggered by Gram-positive bacteria in the cytosol.
Proceedings of the National Academy of Sciences of the United States of America 2004;101(31):11386-91.
2004: Anna I Bakardjiev; Brian A Stacy; Susan J Fisher; Daniel A Portnoy
Listeriosis in the pregnant guinea pig: a model of vertical transmission.
Infection and immunity 2004;72(1):489-97.
2003: Luisa W Cheng; Daniel A Portnoy
Drosophila S2 cells: an alternative infection model for Listeria monocytogenes.
Cellular microbiology 2003;5(12):875-85.
2003: Ian J Glomski; Amy L Decatur; Daniel A Portnoy
Listeria monocytogenes mutants that fail to compartmentalize listerolysin O activity are cytotoxic, avirulent, and unable to evade host extracellular defenses.
Infection and immunity 2003;71(12):6754-65.
2003: Mary O'Riordan; Marlena A Moors; Daniel A Portnoy
Listeria intracellular growth and virulence require host-derived lipoic acid.
Science (New York, N.Y.) 2003;302(5644):462-4.
2003: Laurel L Lenz; Sina Mohammadi; Aimee Geissler; Daniel A Portnoy
SecA2-dependent secretion of autolytic enzymes promotes Listeria monocytogenes pathogenesis.
Proceedings of the National Academy of Sciences of the United States of America 2003;100(21):12432-7.
2003: Victoria Auerbuch; Joseph Loureiro; Frank B Gertler; Julie A Theriot; Daniel A Portnoy
Ena/VASP proteins contribute to Listeria monocytogenes pathogenesis by controlling temporal and spatial persistence of bacterial actin-based motility.
Molecular microbiology 2003;49(5):1361-75.
2002: Christina E Dancz; Andrea Haraga; Daniel A Portnoy; Darren E Higgins
Inducible control of virulence gene expression in Listeria monocytogenes: temporal requirement of listeriolysin O during intracellular infection.
Journal of bacteriology 2002;184(21):5935-45.
2002: Mary O'Riordan; Caroline H Yi; Ramona Gonzales; Kyung-Dall Lee; Daniel A Portnoy
Innate recognition of bacteria by a macrophage cytosolic surveillance pathway.
Proceedings of the National Academy of Sciences of the United States of America 2002;99(21):13861-6.
2002: Daniel A Portnoy; Victoria Auerbuch; Ian J Glomski
The cell biology of Listeria monocytogenes infection: the intersection of bacterial pathogenesis and cell-mediated immunity.
The Journal of cell biology 2002;158(3):409-14.
2002: Laurel L Lenz; Daniel A Portnoy
Identification of a second Listeria secA gene associated with protein secretion and the rough phenotype.
Molecular microbiology 2002;45(4):1043-56.
2002: Mary O'Riordan; Daniel A Portnoy
The host cytosol: front-line or home front?
Trends in microbiology 2002;10(8):361-4.
2002: Peter Lauer; Man Yin Nora Chow; Martin J Loessner; Daniel A Portnoy; Richard Calendar
Construction, characterization, and use of two Listeria monocytogenes site-specific phage integration vectors.
Journal of bacteriology 2002;184(15):4177-86.
2002: Ian J Glomski; Margaret M Gedde; Albert Tsang; Joel A Swanson; Daniel A Portnoy
The Listeria monocytogenes hemolysin has an acidic pH optimum to compartmentalize activity and prevent damage to infected host cells.
The Journal of cell biology 2002;156(6):1029-38.
2002: Victoria Auerbuch; Daniel A Portnoy
Microbial pathogenesis and the eukaryotic cytoskeleton at the American Society for Cell Biology annual meeting.
Cellular microbiology 2002;4(3):187-90.
2001: Peter Lauer; Julie A Theriot; J Skoble; Matt D Welch; Daniel A Portnoy
Systematic mutational analysis of the amino-terminal domain of the Listeria monocytogenes ActA protein reveals novel functions in actin-based motility.
Molecular microbiology 2001;42(5):1163-77.
2001: J Skoble; Victoria Auerbuch; Erin Goley; Matt D Welch; Daniel A Portnoy
Pivotal role of VASP in Arp2/3 complex-mediated actin nucleation, actin branch-formation, and Listeria monocytogenes motility.
The Journal of cell biology 2001;155(1):89-100.
2001: Victoria Auerbuch; Laurel L Lenz; Daniel A Portnoy
Development of a competitive index assay to evaluate the virulence of Listeria monocytogenes actA mutants during primary and secondary infection of mice.
Infection and immunity 2001;69(9):5953-7.
2000: Amy L Decatur; Daniel A Portnoy
A PEST-like sequence in listeriolysin O essential for Listeria monocytogenes pathogenicity.
Science (New York, N.Y.) 2000;290(5493):992-5.
2000: J Skoble; Daniel A Portnoy; Matt D Welch
Three regions within ActA promote Arp2/3 complex-mediated actin nucleation and Listeria monocytogenes motility.
The Journal of cell biology 2000;150(3):527-38.
2000: M M Gedde; Darren E Higgins; L G Tilney; Daniel A Portnoy
Role of listeriolysin O in cell-to-cell spread of Listeria monocytogenes.
Infection and immunity 2000;68(2):999-1003.
1999: Marlena A Moors; Victoria Auerbuch; Daniel A Portnoy
Stability of the Listeria monocytogenes ActA protein in mammalian cells is regulated by the N-end rule pathway.
Cellular microbiology 1999;1(3):249-57.
1999: Darren E Higgins; Nilabh Shastri; Daniel A Portnoy
Delivery of protein to the cytosol of macrophages using Escherichia coli K-12.
Molecular microbiology 1999;31(6):1631-41.
1999: Marlena A Moors; B Levitt; P Youngman; Daniel A Portnoy
Expression of listeriolysin O and ActA by intracellular and extracellular Listeria monocytogenes.
Infection and immunity 1999;67(1):131-9.
1998: R J Thompson; H G Archie Bouwer; Daniel A Portnoy; Fred R Frankel
Pathogenicity and immunogenicity of a Listeria monocytogenes strain that requires D-alanine for growth.
Infection and immunity 1998;66(8):3552-61.
1998: Matt D Welch; J Rosenblatt; J Skoble; Daniel A Portnoy; Timothy J Mitchison
Interaction of human Arp2/3 complex and the Listeria monocytogenes ActA protein in actin filament nucleation.
Science (New York, N.Y.) 1998;281(5373):105-8.
1998: Darren E Higgins; Daniel A Portnoy
Bacterial delivery of DNA evolves.
Nature biotechnology 1998;16(2):138-9.
1997: H Marquis; Howard Goldfine; Daniel A Portnoy
Proteolytic pathways of activation and degradation of a bacterial phospholipase C during intracellular infection by Listeria monocytogenes.
The Journal of cell biology 1997;137(6):1381-92.
1996: G A Smith; J A Theriot; Daniel A Portnoy
The tandem repeat domain in the Listeria monocytogenes ActA protein controls the rate of actin-based motility, the percentage of moving bacteria, and the localization of vasodilator-stimulated phosphoprotein and profilin.
The Journal of cell biology 1996;135(3):647-60.
1996: S Jones; K Preiter; Daniel A Portnoy
Conversion of an extracellular cytolysin into a phagosome-specific lysin which supports the growth of an intracellular pathogen.
Molecular microbiology 1996;21(6):1219-25.
1995: H Marquis; V Doshi; Daniel A Portnoy
The broad-range phospholipase C and a metalloprotease mediate listeriolysin O-independent escape of Listeria monocytogenes from a primary vacuole in human epithelial cells.
Infection and immunity 1995;63(11):4531-4.
1995: G A Smith; H Marquis; S Jones; N C Johnston; Daniel A Portnoy; Howard Goldfine
The two distinct phospholipases C of Listeria monocytogenes have overlapping roles in escape from a vacuole and cell-to-cell spread.
Infection and immunity 1995;63(11):4231-7.
1995: G A Smith; Daniel A Portnoy; J A Theriot
Asymmetric distribution of the Listeria monocytogenes ActA protein is required and sufficient to direct actin-based motility.
Molecular microbiology 1995;17(5):945-51.
1995: Marlena A Moors; Daniel A Portnoy
Identification of bacterial genes that contribute to survival and growth in an intracellular environment.
Trends in microbiology 1995;3(3):83-5.
1994: G Ikonomidis; Yvonne Paterson; F J Kos; Daniel A Portnoy
Delivery of a viral antigen to the class I processing and presentation pathway by Listeria monocytogenes.
The Journal of experimental medicine 1994;180(6):2209-18.
1994: S Jones; Daniel A Portnoy
Characterization of Listeria monocytogenes pathogenesis in a strain expressing perfringolysin O in place of listeriolysin O.
Infection and immunity 1994;62(12):5608-13.
1994: Daniel A Portnoy; S Jones
The cell biology of Listeria monocytogenes infection (escape from a vacuole).
Annals of the New York Academy of Sciences 1994;730():15-25.
1994: Nancy E Freitag; Daniel A Portnoy
Dual promoters of the Listeria monocytogenes prfA transcriptional activator appear essential in vitro but are redundant in vivo.
Molecular microbiology 1994;12(5):845-53.
1994: J A Theriot; J Rosenblatt; Daniel A Portnoy; Pascal J Goldschmidt-Clermont; Timothy J Mitchison
Involvement of profilin in the actin-based motility of L. monocytogenes in cells and in cell-free extracts.
Cell 1994;76(3):505-17.
1994: Daniel A Portnoy; G A Smith; Howard Goldfine
Phospholipases C and the pathogenesis of Listeria.
Brazilian journal of medical and biological research = Revista brasileira de pesquisas médicas e biológicas / Sociedade Brasileira de Biofísica ... [et al.] 1994;27(2):357-61.
1993: R A Brundage; G A Smith; A Camilli; J A Theriot; Daniel A Portnoy
Expression and phosphorylation of the Listeria monocytogenes ActA protein in mammalian cells.
Proceedings of the National Academy of Sciences of the United States of America 1993;90(24):11890-4.
1993: H Marquis; H G Archie Bouwer; David J Hinrichs; Daniel A Portnoy
Intracytoplasmic growth and virulence of Listeria monocytogenes auxotrophic mutants.
Infection and immunity 1993;61(9):3756-60.
1993: G A Smith; Daniel A Portnoy
The role of two phospholipases in the pathogenicity of Listeria monocytogenes.
Infectious agents and disease 1993;2(4):183-5.
1993: Nancy E Freitag; Li Rong; Daniel A Portnoy
Regulation of the prfA transcriptional activator of Listeria monocytogenes: multiple promoter elements contribute to intracellular growth and cell-to-cell spread.
Infection and immunity 1993;61(6):2537-44.
1993: A Camilli; L G Tilney; Daniel A Portnoy
Dual roles of plcA in Listeria monocytogenes pathogenesis.
Molecular microbiology 1993;8(1):143-57.
1992: Daniel A Portnoy; R K Tweten; M Kehoe; J Bielecki
Capacity of listeriolysin O, streptolysin O, and perfringolysin O to mediate growth of Bacillus subtilis within mammalian cells.
Infection and immunity 1992;60(7):2710-7.
1992: H G Archie Bouwer; C S Nelson; Bruce Gibbins; Daniel A Portnoy; David J Hinrichs
Listeriolysin O is a target of the immune response to Listeria monocytogenes.
The Journal of experimental medicine 1992;175(6):1467-71.
1992: J A Theriot; Timothy J Mitchison; L G Tilney; Daniel A Portnoy
The rate of actin-based motility of intracellular Listeria monocytogenes equals the rate of actin polymerization.
Nature 1992;357(6375):257-60.
1992: Ronald A Barry; H G Archie Bouwer; Daniel A Portnoy; David J Hinrichs
Pathogenicity and immunogenicity of Listeria monocytogenes small-plaque mutants defective for intracellular growth and cell-to-cell spread.
Infection and immunity 1992;60(4):1625-32.
1992: Daniel A Portnoy; T Chakraborty; W Goebel; P Cossart
Molecular determinants of Listeria monocytogenes pathogenesis.
Infection and immunity 1992;60(4):1263-7.
1992: Nancy E Freitag; Philip Youngman; Daniel A Portnoy
Transcriptional activation of the Listeria monocytogenes hemolysin gene in Bacillus subtilis.
Journal of bacteriology 1992;174(4):1293-8.
1992: Daniel A Portnoy
Innate immunity to a facultative intracellular bacterial pathogen.
Current opinion in immunology 1992;4(1):20-4.
1991: A Camilli; Howard Goldfine; Daniel A Portnoy
Listeria monocytogenes mutants lacking phosphatidylinositol-specific phospholipase C are avirulent.
The Journal of experimental medicine 1991;173(3):751-4.
1990: Lewis G Tilney; Patricia S Connelly; Daniel A Portnoy
Actin filament nucleation by the bacterial pathogen, Listeria monocytogenes.
The Journal of cell biology 1990;111(6 Pt 2):2979-88.
1990: A N Sun; A Camilli; Daniel A Portnoy
Isolation of Listeria monocytogenes small-plaque mutants defective for intracellular growth and cell-to-cell spread.
Infection and immunity 1990;58(11):3770-8.
1990: Guissou A Dabiri; Jean M Sanger; Daniel A Portnoy; Frederick S Southwick
Listeria monocytogenes moves rapidly through the host-cell cytoplasm by inducing directional actin assembly.
Proceedings of the National Academy of Sciences of the United States of America 1990;87(16):6068-72.
1990: J Bielecki; Philip Youngman; P Connelly; Daniel A Portnoy
Bacillus subtilis expressing a haemolysin gene from Listeria monocytogenes can grow in mammalian cells.
Nature 1990;345(6271):175-6.
1989: Daniel A Portnoy; R D Schreiber; P Connelly; Lewis G Tilney
Gamma interferon limits access of Listeria monocytogenes to the macrophage cytoplasm.
The Journal of experimental medicine 1989;170(6):2141-6.
1989: Lewis G Tilney; Daniel A Portnoy
Actin filaments and the growth, movement, and spread of the intracellular bacterial parasite, Listeria monocytogenes.
The Journal of cell biology 1989;109(4 Pt 1):1597-608.
1989: A Camilli; C R Paynton; Daniel A Portnoy
Intracellular methicillin selection of Listeria monocytogenes mutants unable to replicate in a macrophage cell line.
Proceedings of the National Academy of Sciences of the United States of America 1989;86(14):5522-6.