Kevin Bradley Urdahl, MD, PhD

Kevin Bradley Urdahl, MD, PhD

Infectious Disease Research

Children's Title: Principal Investigator, Supervisor, Center for Global Infectious Disease Research

Academic Title: Professor

Research Center: Center for Global Infectious Disease Research

  • Biography

  • Patient Testimonials

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  • Awards and Honors

    Award Name Award Description Awarded By Award Date
    Member, Society for Pediatric Research 2009
    Research Royalty Fund Pilot Award University of Washington 2007
    Puget Sound Partners in Global Health Pilot Project Award 2006
    Burroughs Wellcome Career Award in the Biological Sciences 2003
    Child Health Research Center Scholar 2003
    Pediatric Immunology Training Grant Fellowship 2002
    Pediatric Scientist Development Program Fellowship 1999
    Bacaner Graduate Medical Research Award, Microbiology (annual award to top Ph.D. graduate) 1993
    J. Thomas Livermore Award, Medical Student Research Award, Hematology 1992
    Graduate School Dissertation Fellowship University of Minnesota 1992
    N.I.H. Immunology Training Grant Fellowship 1991

  • Publications

    Manuscripts in Refereed Journals

    • Stoltzfus CR, Filipek J, Gern BH, Olin BE, Leal JM, Wu Y, Lyons-Cohen MR, Huang JY, Paz-Stoltzfus CL, Plumlee CR, Pöschinger T, Urdahl KB, Perro M, Gerner MY.
      CytoMAP: A Spatial Analysis Toolbox Reveals Features of Myeloid Cell Organization in Lymphoid Tissues.
      Cell Rep., 2020 Apr 21 : 31(3)107523
      https://pmlegacy.ncbi.nlm.nih.gov/pubmed/32320656
    • Delahaye JL, Gern BH, Cohen SB, Plumlee CR, Shafiani S, Gerner MY, Urdahl KB.
      Cutting Edge: Bacillus Calmette-Guérin-Induced T Cells Shape Mycobacterium tuberculosis Infection before Reducing the Bacterial Burden.
      J Immunol, 2019 Aug 15 : 203(4)807-812 PMCID:PMC6684453
      https://www.ncbi.nlm.nih.gov/pubmed/31308091
    • Adams KN, Verma AK, Gopalaswamy R, Adikesavalu H, Singhal DK, Tripathy S, Ranganathan UD, Sherman DR, Urdahl KB, Ramakrishnan L, Hernandez RE.
      Diverse Clinical Isolates of Mycobacterium tuberculosis Develop Macrophage-Induced Rifampin Tolerance.
      J Infect Dis, 2019 Apr 19 : 219(10)1554-1558 PMCID:PMC6473171
      https://www.ncbi.nlm.nih.gov/pubmed/30753612
    • Zhang Q, Chao TC, Patil VS, Qin Y, Tiwari SK, Chiou J, Dobin A, Tsai CM, Li Z, Dang J, Gupta S, Urdahl K, Nizet V, Gingeras TR, Gaulton KJ, Rana TM.
      The long noncoding RNA ROCKI regulates inflammatory gene expression.
      EMBO J, 2019 Apr 15 : 38(8)pii:e100041 PMCID:PMC6463213
      https://www.ncbi.nlm.nih.gov/pubmed/30918008
    • Cohen SB, Urdahl KB.
      Going beyond gamma for TB protection.
      Nat Microbiol, 2018 Nov. : 3(11)1194-1195
      https://www.ncbi.nlm.nih.gov/pubmed/30356152
    • Cohen SB, Gern BH, Delahaye JL, Adams KN, Plumlee CR, Winkler JK, Sherman DR, Gerner MY, Urdahl KB.
      Alveolar Macrophages Provide an Early Mycobacterium tuberculosis Niche and Initiate Dissemination.
      Cell Host Microbe, 2018 Sep 12 : 24(3)439-446.e4 PMCID:PMC6152889
      https://www.ncbi.nlm.nih.gov/pubmed/30146391
    • Lindenstrøm T, Moguche A, Damborg M, Agger EM, Urdahl K, Andersen P.
      T Cells Primed by Live Mycobacteria Versus a Tuberculosis Subunit Vaccine Exhibit Distinct Functional Properties.
      EBioMedicine, 2018 Jan. : 2727-39 PMCID:PMC5828549
      https://www.ncbi.nlm.nih.gov/pubmed/29249639
    • Moguche AO, Musvosvi M, Penn-Nicholson A, Plumlee CR, Mearns H, Geldenhuys H, Smit E, Abrahams D, Rozot V, Dintwe O, Hoff ST, Kromann I, Ruhwald M, Bang P, Larson RP, Shafiani S, Ma S, Sherman DR, Sette A, Lindestam Arlehamn CS, McKinney DM, Maecker H, Hanekom WA, Hatherill M, Andersen P, Scriba TJ, Urdahl KB.
      Antigen Availability Shapes T Cell Differentiation and Function during Tuberculosis.
      Cell Host Microbe, 2017 Jun 14 : 21(6)695-706.e5 PMCID:PMC5533182
      https://www.ncbi.nlm.nih.gov/pubmed/28618268
    • Woodworth JS, Cohen SB, Moguche AO, Plumlee CR, Agger EM, Urdahl KB, Andersen P.
      Subunit vaccine H56/CAF01 induces a population of circulating CD4 T cells that traffic into the Mycobacterium tuberculosis-infected lung.
      Mucosal Immunol, 2017 March : 10(2)555-564 PMCID:PMC5325828
      https://www.ncbi.nlm.nih.gov/pubmed/27554293
    • Rothchild AC, Sissons JR, Shafiani S, Plaisier C, Min D, Mai D, Gilchrist M, Peschon J, Larson RP, Bergthaler A, Baliga NS, Urdahl KB, Aderem A.
      MiR-155-regulated molecular network orchestrates cell fate in the innate and adaptive immune response to Mycobacterium tuberculosis.
      Proc Natl Acad Sci U S A, 2016 Oct 11 : 113(41)E6172-E6181 PMCID:PMC5068277
      https://www.ncbi.nlm.nih.gov/pubmed/27681624
    • Levitte S, Adams KN, Berg RD, Cosma CL, Urdahl KB, Ramakrishnan L.
      Mycobacterial Acid Tolerance Enables Phagolysosomal Survival and Establishment of Tuberculous Infection In Vivo.
      Cell Host Microbe, 2016 Aug 10 : 20(2)250-8 PMCID:PMC4985559
      https://www.ncbi.nlm.nih.gov/pubmed/27512905
    • Di Paolo NC, Shafiani S, Day T, Papayannopoulou T, Russell DW, Iwakura Y, Sherman D, Urdahl K, Shayakhmetov DM.
      Interdependence between Interleukin-1 and Tumor Necrosis Factor Regulates TNF-Dependent Control of Mycobacterium tuberculosis Infection.
      Immunity, 2015 Dec 15 : 43(6)1125-36 PMCID:PMC4685953
      https://www.ncbi.nlm.nih.gov/pubmed/26682985
    • Moguche AO, Shafiani S, Clemons C, Larson RP, Dinh C, Higdon LE, Cambier CJ, Sissons JR, Gallegos AM, Fink PJ, Urdahl KB.
      ICOS and Bcl6-dependent pathways maintain a CD4 T cell population with memory-like properties during tuberculosis.
      J Exp Med, 2015 May 4 : 212(5)715-28 PMCID:PMC4419347
      https://www.ncbi.nlm.nih.gov/pubmed/25918344
    • Cambier CJ, Takaki KK, Larson RP, Hernandez RE, Tobin DM, Urdahl KB, Cosma CL, Ramakrishnan L.
      Mycobacteria manipulate macrophage recruitment through coordinated use of membrane lipids.
      Nature, 2014 Jan 9 : 505(7482)218-22 PMCID:PMC3961847
      https://www.ncbi.nlm.nih.gov/pubmed/24336213
    • Shafiani S, Dinh C, Ertelt JM, Moguche AO, Siddiqui I, Smigiel KS, Sharma P, Campbell DJ, Way SS, Urdahl KB.
      Pathogen-specific Treg cells expand early during Mycobacterium tuberculosis infection but are later eliminated in response to Interleukin-12.
      Immunity, 2013 Jun 27 : 38(6)1261-70 PMCID:PMC3827956
      https://www.ncbi.nlm.nih.gov/pubmed/23791647
    • Reiley WW, Shafiani S, Wittmer ST, Tucker-Heard G, Moon JJ, Jenkins MK, Urdahl KB, Winslow GM, Woodland DL.
      Distinct functions of antigen-specific CD4 T cells during murine Mycobacterium tuberculosis infection.
      Proc Natl Acad Sci U S A, 2010 Nov 9 : 107(45)19408-13 PMCID:PMC2984157
      https://www.ncbi.nlm.nih.gov/pubmed/20962277
    • Curtis MM, Rowell E, Shafiani S, Negash A, Urdahl KB, Wilson CB, Way SS.
      Fidelity of pathogen-specific CD4+ T cells to the Th1 lineage is controlled by exogenous cytokines, interferon-gamma expression, and pathogen lifestyle.
      Cell Host Microbe, 2010 Aug 19 : 8(2)163-73 PMCID:PMC2923648
      https://www.ncbi.nlm.nih.gov/pubmed/20709293
    • Shafiani S, Tucker-Heard G, Kariyone A, Takatsu K, Urdahl KB.
      Pathogen-specific regulatory T cells delay the arrival of effector T cells in the lung during early tuberculosis.
      J Exp Med, 2010 Jul 5 : 207(7)1409-20 PMCID:PMC2901066
      https://www.ncbi.nlm.nih.gov/pubmed/20547826
    • Koch MA, Tucker-Heard G, Perdue NR, Killebrew JR, Urdahl KB, Campbell DJ.
      The transcription factor T-bet controls regulatory T cell homeostasis and function during type 1 inflammation.
      Nat Immunol, 2009 June : 10(6)595-602 PMCID:PMC2712126
      https://www.ncbi.nlm.nih.gov/pubmed/19412181
    • Scott-Browne JP, Shafiani S, Tucker-Heard G, Ishida-Tsubota K, Fontenot JD, Rudensky AY, Bevan MJ, Urdahl KB.
      Expansion and function of Foxp3-expressing T regulatory cells during tuberculosis.
      J Exp Med, 2007 Sep 3 : 204(9)2159-69 PMCID:PMC2118702
      https://www.ncbi.nlm.nih.gov/pubmed/17709423
    • Urdahl KB, Liggitt D, Bevan MJ.
      CD8+ T cells accumulate in the lungs of Mycobacterium tuberculosis-infected Kb-/-Db-/- mice, but provide minimal protection.
      J Immunol, 2003 Feb 15 : 170(4)1987-94
      https://www.ncbi.nlm.nih.gov/pubmed/12574368
    • Urdahl KB, Sun JC, Bevan MJ.
      Positive selection of MHC class Ib-restricted CD8(+) T cells on hematopoietic cells.
      Nat Immunol, 2002 Aug. : 3(8)772-9 PMCID:PMC2782383
      https://www.ncbi.nlm.nih.gov/pubmed/12089507
    • Qin X, Urdahl KB.
      PCR and sequencing of independent genetic targets for the diagnosis of culture negative bacterial endocarditis.
      Diagn Microbiol Infect Dis, 2001 Aug. : 40(4)145-9
      https://www.ncbi.nlm.nih.gov/pubmed/11576785
    • Shi YP, Nahlen BL, Kariuki S, Urdahl KB, McElroy PD, Roberts JM, Lal AA.
      Fcgamma receptor IIa (CD32) polymorphism is associated with protection of infants against high-density Plasmodium falciparum infection. VII. Asembo Bay Cohort Project.
      J Infect Dis, 2001 Jul 1 : 184(1)107-11
      https://www.ncbi.nlm.nih.gov/pubmed/11398118
    • Chou TD, Gibran NS, Urdahl K, Lin EY, Heimbach DM, Engrav LH.
      Methemoglobinemia secondary to topical silver nitrate therapy--a case report.
      Burns, 1999 Sept. : 25(6)549-52
      https://www.ncbi.nlm.nih.gov/pubmed/10498367
    • Urdahl KB, Mathews JD, Currie B.
      Anti-streptokinase antibodies and streptokinase resistance in an Aboriginal population in northern Australia.
      Aust N Z J Med, 1996 Feb. : 26(1)49-53
      https://www.ncbi.nlm.nih.gov/pubmed/8775528
    • Urdahl KB, Pardoll DM, Jenkins MK.
      Cyclosporin A inhibits positive selection and delays negative selection in alpha beta TCR transgenic mice.
      J Immunol, 1994 Mar 15 : 152(6)2853-9
      https://www.ncbi.nlm.nih.gov/pubmed/8144886
    • Urdahl KB, Pardoll DM, Jenkins MK.
      Self-reactive T cells are present in the peripheral lymphoid tissues of cyclosporin A-treated mice.
      Int Immunol, 1992 Dec. : 4(12)1341-9
      https://www.ncbi.nlm.nih.gov/pubmed/1286060
    • Norton SD, Zuckerman L, Urdahl KB, Shefner R, Miller J, Jenkins MK.
      The CD28 ligand, B7, enhances IL-2 production by providing a costimulatory signal to T cells.
      J Immunol, 1992 Sep 1 : 149(5)1556-61
      https://www.ncbi.nlm.nih.gov/pubmed/1380533
    • Haase AT, Jenkins MK, Peng H, Urdahl K.
      nef-naf nexus?
      Curr Biol, 1992 March : 2(3)130-2
      https://www.ncbi.nlm.nih.gov/pubmed/15335987
    • DeSilva DR, Urdahl KB, Jenkins MK.
      Clonal anergy is induced in vitro by T cell receptor occupancy in the absence of proliferation.
      J Immunol, 1991 Nov 15 : 147(10)3261-7
      https://www.ncbi.nlm.nih.gov/pubmed/1658142
    • Jenkins MK, Taylor PS, Norton SD, Urdahl KB.
      CD28 delivers a costimulatory signal involved in antigen-specific IL-2 production by human T cells.
      J Immunol, 1991 Oct 15 : 147(8)2461-6
      https://www.ncbi.nlm.nih.gov/pubmed/1717561
    • O'Shea JJ, Urdahl KB, Luong HT, Chused TM, Samelson LE, Klausner RD.
      Aluminum fluoride induces phosphatidylinositol turnover, elevation of cytoplasmic free calcium, and phosphorylation of the T cell antigen receptor in murine T cells.
      J Immunol, 1987 Nov 15 : 139(10)3463-9
      https://www.ncbi.nlm.nih.gov/pubmed/2824607
    • Samelson LE, O'Shea JJ, Luong H, Ross P, Urdahl KB, Klausner RD, Bluestone J.
      T cell antigen receptor phosphorylation induced by an anti-receptor antibody.
      J Immunol, 1987 Oct 15 : 139(8)2708-14
      https://www.ncbi.nlm.nih.gov/pubmed/2443570

    Book Chapters

    • Larson RP, Shafiani S, Urdahl KB.
      Foxp3(+) regulatory T cells in tuberculosis.
      The New Paradigm of Immunity to Tuberculosis, Adv Exp Med Biol, Springer, New York, 2013 : 783165-80
      https://www.ncbi.nlm.nih.gov/pubmed/23468109
    • Urdahl KB, Jenkins MK
      Antigen-presenting cell regulation of T cell activation.
      Handbook of B and T lymphocytes, Snow EC (ed.), Academic Press, San Diego, 1994 : 143-53
    • Urdahl KB, Jenkins MK, Norton SD.
      Accessory cell-derived costimulatory signals regulate T cell proliferation.
      Ann N Y Acad Sci, 1991 Dec 30 : 63633-42
      https://www.ncbi.nlm.nih.gov/pubmed/1665322
    • Jenkins MK, Mueller D, Schwartz RH, Carding S, Bottomley K, Stadecker MJ, Urdahl KB, Norton SD.
      Induction and maintenance of anergy in mature T cells.
      Mechanisms of lymphocyte activation and immune regulation III: Developmental biology of lymphocytes, Gupta S, Paul W, Cooper M, and Rotherberg E (eds)., Plenum Press, New York, 1991 : 292167-76
      https://www.ncbi.nlm.nih.gov/pubmed/1950767

    Other Publications

    • Andersen P, Urdahl KB.
      TB vaccines; promoting rapid and durable protection in the lung.
      Curr Opin Immunol, 2015 Aug. : 3555-62. Review. PMCID:PMC4641675
      https://www.ncbi.nlm.nih.gov/pubmed/26113434
    • Urdahl KB.
      Understanding and overcoming the barriers to T cell-mediated immunity against tuberculosis.
      Semin Immunol, 2014 Dec. : 26(6)578-87. Review. PMCID:PMC4314386
      https://www.ncbi.nlm.nih.gov/pubmed/25453230
    • Vanden Driessche K, Persson A, Marais BJ, Fink PJ, Urdahl KB.
      Immune vulnerability of infants to tuberculosis.
      Clin Dev Immunol, 2013 Feb. : 2013781320. Review. PMCID:PMC3666431
      https://www.ncbi.nlm.nih.gov/pubmed/23762096
    • Urdahl KB, Shafiani S, Ernst JD.
      Initiation and regulation of T-cell responses in tuberculosis.
      Mucosal Immunol, 2011 May : 4(3)288-93. Review. PMCID:PMC3206635
      https://www.ncbi.nlm.nih.gov/pubmed/21451503

  • Presentations

    Presentation Title Event Location Date
    TGFß restricts T cell-mediated immunity in the TB granuloma InterCenter Discovery Lunch Seattle Children's Research Institute, Seattle WA 2019
    T cell mediated immunity in the tuberculous granuloma Cambridge Immunology Network Seminar MRC Laboratory of Molecular Biology, University of Cambridge 2017
    Basic Immunology and Developing an Effective TB Vaccine 3rd Annual Collaboration for TB Vaccine Discovery Meeting Bill and Melinda Gates Foundation, Seattle WA 2017
    Of Mice and Men: Tuberculosis Fiction or a Model of Human Disease? Microbiology and Immunobiology Seminar Harvard University 2017
    Of Mice and Men: Tuberculosis Fiction or a Model of Human Disease? Microbiology and Immunology Seminar University of California, Berkeley 2017
    Of Mice and Men: Tuberculosis Fiction or a Model of Human Disease? Microbiology and Immunology Seminar Albert Einstein College of Medicine, Bronx NY 2016
    Of Mice and Men: Tuberculosis Fiction or a Model of Human Disease? Division of Immunology and Pathogenesis, University of California, Berkeley 2016
    Of Mice and Men: Tuberculosis Fiction or a Model of Human Disease? Microbiology and Immunology Seminar Albert Einstein College of Medicine, Bronx NY 2016
    Update on our understanding of conventional T cells and immunity against TB 2nd Annual Collaboration for TB Vaccine Discovery Meeting Bill and Melinda Gates Foundation, Seattle WA 2016
    Mucosal immunity to bacterial pathogens Block Symposium, 103st Annual Meeting American Association of Immunologists, Seattle WA 2016
    The complex relationship between effector T cell specificity, localization, and function during pulmonary tuberculosis Copenhagen, Denmark 2015
    Understanding and overcoming the barriers to T cell mediated immunity against tuberculosis Seminar Series University of Iowa, Iowa City IA 2015
    Understanding and overcoming the barriers to T cell mediated immunity against tuberculosis Seminar Series Seattle Biomedical Research Institute, Seattle WA 2014
    Regulatory T cells in tuberculosis Seminar Series University of Iowa, Iowa City IA 2014
    The complex relationship between effector T cell specificity, location, and function during pulmonary TB Seminar Series Trudeau Institute, Saranac Lake NY 2014
    Immunology of the Airway Block Symposium, 101st Annual Meeting American Association of Immunologists, Pittsburgh PA 2014
    The complex relationship between effector T cell specificity, location, and function during pulmonary TB. Cold Spring Harbor Laboratory Meeting: Harnessing Immunity to Prevent and Treat Disease Cold Spring Harbor, NY 2013
    T cell responses to Mycobacterium tuberculosis: Why are they so slow and how do they persist? Department of Pathobiology Seminar Series University of Washington 2012
    Immune responses that restrict and promote long lasting protection against Mycobacterium tuberculosis: Lessons learned in mice. Vaccine and Infectious Disease Division Faculty Seminar Series Fred Hutchinson Cancer Research Center, Seattle WA 2011
    Pathogen-specific regulatory T cells and the slow-motion response to Mycobacterium tuberculosis infection. 15th Annual Amgen Joint Research Symposium Seattle, WA 2011
    Pathogen-specific regulatory T cells and the slow-motion response to Mycobacterium tuberculosis infection. Microbiology, Immunology and Cancer Biology, Seminar Series University of Minnesota, Minneapolis, MN 2011
    Regulation and memory in tuberculosis. Plenary Session, Keystone Symposium, Tuberculosis: Immunology, Cell Biology, and Novel Vaccination Strategies Vancouver, British Columbia 2011
    Pathogen-specific regulatory T cells and the slow-motion response to Mycobacterium tuberculosis infection. Annual Retreat, Department of Immunology University of Washington, Seattle, WA 2010
    Pathogen-specific regulatory T cells impair protective immunity in early Mycobacterium tuberculosis infection. Infectious Disease Immunology Symposium Seattle Biomedical Research Institute, Seattle, WA 2009
    Pathogen-specific regulatory T cells impair protective immunity in early Mycobacterium tuberculosis infection. Center for Lung Biology, University of Washington, Seattle, WA. 2009
    Pathogen-specific regulatory T cells impair protective immunity against tuberculosis. Laboratory of Parasitic Diseases Seminar Series NIH, NIAID, Bethesda, MD 2008
    Foxp3-expressing regulatory T cells in tuberculosis. Major Symposium, 95th Annual Meeting American Association of Immunologists, San Diego, CA 2008
    Location and function of Foxp3-expressing T regulatory cells in tuberculosis. Plenary Session, Keystone Symposium, Host-Pathogen Standoff: Persistent and Latent Infection Keystone, Denver, CO 2006
    Diagnosis, treatment, and prevention of tuberculosis in children: Limitations of current modalities, and possibilities for the future. Grand Rounds Seattle Children's Hospital 2005
    Foxp3-expressing T regulatory cells in tuberculosis. 40th Annual U.S.-Japan Cooperative Medical Science Program’s Tuberculosis and Leprosy Conference Seattle, WA 2005
    The role of MHC class I molecules in immunity against tuberculosis. International Training and Research in Emerging Infections Diseases, Intracellular Pathogens New Delhi, India 2005
    T cells and TB: Lessons from the Mouse Model. Department of Pathobiology Seminar Series University of Washington 2005
    The role of MHC class I molecules in immunity against tuberculosis. Departments of Microbiology and Pediatrics University of Minnesota 2003
    The role of MHC class I molecules in immunity against tuberculosis. Seattle Biomedical Research Institute, Seattle, WA 2003

  • Research Funding

    Grant Title Grantor Amount Award Date
    Characterization of Recent Thymic Emigrants - R01AI064318 (PI: Urdahl) NIH/NIAID $470,750/year March 2019 - Feb. 2020
    TOLLIP Deficiency, Immune Dysregulation, and Tuberculosis Susceptibility - 1R01AI136921-01 (PI: Shah) Role: Co-Investigator NIH/NIAID $50,000/year March 2018 - Feb. 2023
    Omics for TB: Response to Infection and Treatment - 1U19AI35976-01 (PI: Aderem/Sherman) Role: Co-Investigator NIH/NIAID $285,652/year Feb. 2018 - Jan. 2023
    A dual function TB subunit vaccine designed for non-interference with BCG and post-exposure activity - 1R01AI135721-01 (PI: Andersen) Role: Co-Investigator NIH/NIAID $15,355/year Dec. 2017 - Nov. 2022
    Eliciting lung-localized CD4 T cell responses against Mycobacterium tuberculosis - 1R01AI134246-01 (PI: Urdahl) NIH/NIAID $206,932/year Aug. 2017 - 2022
    HIV Integration-Mediated Modulation of Immune Regulation in HPV-Associated Cancers - 5R01CA206466-02 (PI: Frenkel) Role: Co-Investigator NIH/NCI $2007/year Sept. 2016 - Aug. 2021

Overview

Clinical Interests

Tuberculosis; Protective immunity against tuberculosis; Tuberculosis vaccines

Research Description

My laboratory is applying recently-developed immunologic tools to the study of tuberculosis in order to gain a better understanding of T lymphocyte-mediated mechanisms that promote protection against tuberculosis and also mechanisms that suppress immunity and help facilitate persistence of the pathogen. The overall goal of this research is to identify strategies that could contribute to the development of an effective vaccine. 

Recently, my laboratory defined a role for a suppressive subset of T lymphocytes that dampens immunity and prevents the immune system from effectively eradicating the bacteria that causes tuberculosis.

Research Focus Area

Host-Pathogen Interaction, Host: Pathogen Interaction, Infectious Disease, Tuberculosis