Allen Ryan

TitleRecall Faculty
InstitutionUniversity of California San Diego
Address9500 Gilman Drive #0666
La Jolla CA 92093
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    Research Overview

    Biology of the inner and middle ears, including hair cell gene and fate regulation, drug and noise damage to hearing, inner ear neuronal targeting, pathogenesis and recovery mechanisms in otitis media, and drug delivery to the middle and inner ears.

    Research Topics

    • Cell Biology

    • Development

    • Drug Delivery

    • Genomics

    • Immunology

    • Infectious Disease

    • Inflammation

    • Injury

    • Innate Immunity

    • Molecular Biology

    • Neuroscience

    • Regeneration/Stem Cells

    • Structural Biology

    • Translational Research

    Current Project Titles

    • Recovery from blast-induced hearing loss (DoD)

    • A biological interface for the cochlear implant (VA)

    • Middle ear responses in otitis media (NIH)

    • Mechanisms of persistence and recovery in otitis media (NIH)

    • New methods for the treatment of hearing and balance disorders (Pending)


    My laboratory studies the basic biology of hearing. A major focus of the group is the sensory hair cells of the cochlea. This includes studies of gene regulation in these cells, as well as fate determination by combinatorial transcription factor coding. We also study cellular mechanisms of hair cell damage and death induced by ototoxic drugs, noise trauma and excitotoxicity. A second area of interest is the two types of afferent neurons in the cochlea. In particular the type II neurons that innervate outer hair cells are poorly understood with respect to their function and the regulation of their complex pathfinding to their target hair cells. We also study pathogenesis and recovery of the middle ear mucosa during otitis media, with a focus on cell phenotype determination and innate immunity. Finally, we are interested in novel methods for drug delivery to the middle and inner ears.

    Collapse Research 
    Collapse Research Activities and Funding
    A Biological Interface for Auditory Rehabilitation with a Cochlear Implant
    VA I01RX000977Oct 1, 2013 - Sep 30, 2016
    Role: Principal Investigator
    Basic Mechanisms in Hearing Loss of Cochlear Origin
    VA I01BX001205Oct 1, 2012 - Sep 30, 2020
    Role: Principal Investigator
    Innovative therapy for diseases of the middle ear
    NIH/NIDCD R01DC012595Jul 1, 2012 - Jun 30, 2017
    Role: Principal Investigator
    The 8th Conference on the Molecular Biology of Hearing &Deafness
    NIH/NIDCD R13DC012023Jul 4, 2011 - Jun 30, 2014
    Role: Principal Investigator
    Conference on Molecular Biology of Hearing and Deafness
    NIH/NIDCD R13DC010087Mar 1, 2009 - Feb 28, 2010
    Role: Principal Investigator
    Conference on the Molecular Biology of Hearing/Deafness
    NIH/NIDCD R13DC007068Sep 1, 2004 - Aug 31, 2005
    Role: Principal Investigator
    NIH/NIDCD R13DC005016Sep 1, 2001 - Aug 31, 2002
    Role: Principal Investigator
    NIH/NIDCD R13DC003804Aug 1, 1998 - Jan 31, 1999
    Role: Principal Investigator
    NIH/NIDCD R13DC002735Jul 1, 1995 - Jun 30, 1996
    Role: Principal Investigator
    NIH/NIDCD R13DC001516May 1, 1992 - Apr 30, 1993
    Role: Principal Investigator
    Otolaryngology Training in Immunology, Virology and Molecular Biology
    NIH/NIDCD T32DC000028Jul 10, 1989 - Jun 30, 2021
    Role: Principal Investigator
    NIH/NIDCD R01DC000139Jul 1, 1979 - Jan 31, 2009
    Role: Principal Investigator
    NIH/NINDS R01NS014945Jul 1, 1979 - Mar 31, 1989
    Role: Principal Investigator
    Middle Ear Response in Otitis Media
    NIH/NIDCD R01DC000129Apr 1, 1978 - Jul 31, 2022
    Role: Principal Investigator
    NIH/NINDS R01NS014389Apr 1, 1978 - Mar 31, 1990
    Role: Principal Investigator

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    Collapse Bibliographic 
    Collapse Publications
    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Researchers can login to make corrections and additions, or contact us for help.
    List All   |   Timeline
    1. Santos-Cortez RLP, Chiong CM, Frank DN, Ryan AF, Giese APJ, Bootpetch Roberts T, Daly KA, Steritz MJ, Szeremeta W, Pedro M, Pine H, Yarza TKL, Scholes MA, Llanes EGDV, Yousaf S, Friedman N, Tantoco MLC, Wine TM, Labra PJ, Benoit J, Ruiz AG, de la Cruz RAR, Greenlee C, Yousaf A, Cardwell J, Nonato RMA, Ray D, Ong KMC, So E, Robertson CE, Dinwiddie J, Lagrana-Villagracia SM, Gubbels SP, Shaikh RS, Cass SP, Einarsdottir E, Lee NR, Schwartz DA, Gloria-Cruz TLI, Bamshad MJ, Yang IV, Kere J, Abes GT, Prager JD, Riazuddin S, Chan AL, Yoon PJ, Nickerson DA, Cutiongco-de la Paz EM, Streubel SO, Reyes-Quintos MRT, Jenkins HA, Mattila P, Chan KH, Mohlke KL, Leal SM, Hafrén L, Chonmaitree T, Sale MM, Ahmed ZM. FUT2 Variants Confer Susceptibility to Familial Otitis Media. Am J Hum Genet. 2018 Nov 01; 103(5):679-690. PMID: 30401457.
      View in: PubMed
    2. Kurabi A, Schaerer D, Noack V, Bernhardt M, Pak K, Alexander T, Husseman J, Nguyen Q, Harris JP, Ryan AF. Active Transport of Peptides Across the Intact Human Tympanic Membrane. Sci Rep. 2018 Aug 07; 8(1):11815. PMID: 30087425.
      View in: PubMed
    3. Lim HW, Pak K, Ryan AF, Kurabi A. Screening Mammalian Cochlear Hair Cells to Identify Critical Processes in Aminoglycoside-Mediated Damage. Front Cell Neurosci. 2018; 12:179. PMID: 30013464.
      View in: PubMed
    4. Hur DG, Kurabi A, Ryan AF. Screening antioxidants for the protection of cochlear sensory cells. Neural Regen Res. 2018 Jan; 13(1):62-64. PMID: 29451206.
      View in: PubMed
    5. Ryals M, Pak K, Jalota R, Kurabi A, Ryan AF. A kinase inhibitor library screen identifies novel enzymes involved in ototoxic damage to the murine organ of Corti. PLoS One. 2017; 12(10):e0186001. PMID: 29049311.
      View in: PubMed
    6. Deniffel D, Nuyen B, Pak K, Suzukawa K, Hung J, Kurabi A, Wasserman SI, Ryan AF. Otitis Media and Nasopharyngeal Colonization in ccl3-/- Mice. Infect Immun. 2017 11; 85(11). PMID: 28847849.
      View in: PubMed
    7. Noack V, Pak K, Jalota R, Kurabi A, Ryan AF. An Antioxidant Screen Identifies Candidates for Protection of Cochlear Hair Cells from Gentamicin Toxicity. Front Cell Neurosci. 2017; 11:242. PMID: 28867994.
      View in: PubMed
    8. Kurabi A, Schaerer D, Chang L, Pak K, Ryan AF. Optimisation of peptides that actively cross the tympanic membrane by random amino acid extension: a phage display study. J Drug Target. 2018 Feb; 26(2):127-134. PMID: 28658990.
      View in: PubMed
    9. Kyd JM, Hotomi M, Kono M, Kurabi A, Pichichero M, Ryan A, Swords WE, Thornton R. Panel 5: Immunology. Otolaryngol Head Neck Surg. 2017 Apr; 156(4_suppl):S63-S75. PMID: 28372530.
      View in: PubMed
    10. Kurabi A, Beasley KA, Chang L, McCann J, Pak K, Ryan AF. Peptides actively transported across the tympanic membrane: Functional and structural properties. PLoS One. 2017; 12(2):e0172158. PMID: 28234923.
      View in: PubMed
    11. Hickox AE, Wong AC, Pak K, Strojny C, Ramirez M, Yates JR, Ryan AF, Savas JN. Global Analysis of Protein Expression of Inner Ear Hair Cells. J Neurosci. 2017 Feb 01; 37(5):1320-1339. PMID: 28039372.
      View in: PubMed
    12. Kurabi A, Keithley EM, Housley GD, Ryan AF, Wong AC. Cellular mechanisms of noise-induced hearing loss. Hear Res. 2017 06; 349:129-137. PMID: 27916698.
      View in: PubMed
    13. Cho CG, Pak K, Webster N, Kurabi A, Ryan AF. Both canonical and non-canonical NF-?B activation contribute to the proliferative response of the middle ear mucosa during bacterial infection. Innate Immun. 2016 11; 22(8):626-634. PMID: 27655045.
      View in: PubMed
    14. Masuda M, Li Y, Pak K, Chavez E, Mullen L, Ryan AF. The Promoter and Multiple Enhancers of the pou4f3 Gene Regulate Expression in Inner Ear Hair Cells. Mol Neurobiol. 2017 Sep; 54(7):5414-5426. PMID: 27592349.
      View in: PubMed
    15. Ryan AF, Kujawa SG, Hammill T, Le Prell C, Kil J. Temporary and Permanent Noise-induced Threshold Shifts: A Review of Basic and Clinical Observations. Otol Neurotol. 2016 09; 37(8):e271-5. PMID: 27518135; PMCID: PMC4988324 [Available on 09/01/17].
    16. Chan P, Ho K, Ryan AF. Impulse Noise Injury Model. Mil Med. 2016 05; 181(5 Suppl):59-69. PMID: 27168554.
      View in: PubMed
    17. Kurabi A, Pak KK, Bernhardt M, Baird A, Ryan AF. Discovery of a Biological Mechanism of Active Transport through the Tympanic Membrane to the Middle Ear. Sci Rep. 2016 Mar 07; 6:22663. PMID: 26946957; PMCID: PMC4780071.
    18. Kurabi A, Pak K, Ryan AF, Wasserman SI. Innate Immunity: Orchestrating Inflammation and Resolution of Otitis Media. Curr Allergy Asthma Rep. 2016 Jan; 16(1):6. PMID: 26732809.
      View in: PubMed
    19. Ryan AF, Dobie RA, Dubno JR, Lonsbury-Martin BL. Tinnitus: Research supported by the Tinnitus Research Consortium. Hear Res. 2016 04; 334:1. PMID: 26739791.
      View in: PubMed
    20. Harrop-Jones A, Wang X, Fernandez R, Dellamary L, Ryan AF, LeBel C, Piu F. The Sustained-Exposure Dexamethasone Formulation OTO-104 Offers Effective Protection against Noise-Induced Hearing Loss. Audiol Neurootol. 2016; 21(1):12-21. PMID: 26655654.
      View in: PubMed
    21. Morton-Jones RT, Vlajkovic SM, Thorne PR, Cockayne DA, Ryan AF, Housley GD. Properties of ATP-gated ion channels assembled from P2X2 subunits in mouse cochlear Reissner's membrane epithelial cells. Purinergic Signal. 2015 Dec; 11(4):551-60. PMID: 26424615; PMCID: PMC4648790.
    22. Froud KE, Wong AC, Cederholm JM, Klugmann M, Sandow SL, Julien JP, Ryan AF, Housley GD. Type II spiral ganglion afferent neurons drive medial olivocochlear reflex suppression of the cochlear amplifier. Nat Commun. 2015 May 12; 6:7115. PMID: 25965946; PMCID: PMC4432632.
    23. Wong AC, Ryan AF. Mechanisms of sensorineural cell damage, death and survival in the cochlea. Front Aging Neurosci. 2015; 7:58. PMID: 25954196; PMCID: PMC4404918.
    24. Hernandez M, Leichtle A, Pak K, Webster NJ, Wasserman SI, Ryan AF. The transcriptome of a complete episode of acute otitis media. BMC Genomics. 2015 Apr 03; 16:259. PMID: 25888408; PMCID: PMC4394589.
    25. Leichtle A, Klenke C, Ebmeyer J, Daerr M, Bruchhage KL, Hoffmann AS, Ryan AF, Wollenberg B, Sudhoff H. NOD-Like Receptor Signaling in Cholesteatoma. Biomed Res Int. 2015; 2015:408169. PMID: 25922834; PMCID: PMC4398947.
    26. Ikeda R, Pak K, Chavez E, Ryan AF. Transcription factors with conserved binding sites near ATOH1 on the POU4F3 gene enhance the induction of cochlear hair cells. Mol Neurobiol. 2015 Apr; 51(2):672-84. PMID: 25015561; PMCID: PMC4291311.
    27. Brand Y, Levano S, Radojevic V, Naldi AM, Setz C, Ryan AF, Pak K, Hemmings BA, Bodmer D. All Akt isoforms (Akt1, Akt2, Akt3) are involved in normal hearing, but only Akt2 and Akt3 are involved in auditory hair cell survival in the mammalian inner ear. PLoS One. 2015; 10(3):e0121599. PMID: 25811375; PMCID: PMC4374771.
    28. Ryan AF, Ikeda R, Masuda M. The regulation of gene expression in hair cells. Hear Res. 2015 Nov; 329:33-40. PMID: 25616095; PMCID: PMC4510037 [Available on 11/01/16].
    29. Brand Y, Sung M, Pak K, Chavez E, Wei E, Radojevic V, Bodmer D, Ryan AF. Neural cell adhesion molecule NrCAM is expressed in the mammalian inner ear and modulates spiral ganglion neurite outgrowth in an in vitro alternate choice assay. J Mol Neurosci. 2015 Apr; 55(4):836-44. PMID: 25407819; PMCID: PMC4355056.
    30. Palmisano MG, Bremner SN, Hornberger TA, Meyer GA, Domenighetti AA, Shah SB, Kiss B, Kellermayer M, Ryan AF, Lieber RL. Skeletal muscle intermediate filaments form a stress-transmitting and stress-signaling network. J Cell Sci. 2015 Jan 15; 128(2):219-24. PMID: 25413344; PMCID: PMC4294770.
    31. Yao W, Frie M, Pan J, Pak K, Webster N, Wasserman SI, Ryan AF. C-Jun N-terminal kinase (JNK) isoforms play differing roles in otitis media. BMC Immunol. 2014 Oct 14; 15:46. PMID: 25311344; PMCID: PMC4200133.
    32. Suzukawa K, Tomlin J, Pak K, Chavez E, Kurabi A, Baird A, Wasserman SI, Ryan AF. A mouse model of otitis media identifies HB-EGF as a mediator of inflammation-induced mucosal proliferation. PLoS One. 2014; 9(7):e102739. PMID: 25033458; PMCID: PMC4102546.
    33. Sung M, Wei E, Chavez E, Jain N, Levano S, Binkert L, Ramseier A, Setz C, Bodmer D, Ryan AF, Brand Y. Inhibition of MMP-2 but not MMP-9 influences inner ear spiral ganglion neurons in vitro. Cell Mol Neurobiol. 2014 Oct; 34(7):1011-21. PMID: 24935409; PMCID: PMC4386629.
    34. Kurabi A, Lee J, Wong C, Pak K, Hoffman HM, Ryan AF, Wasserman SI. The inflammasome adaptor ASC contributes to multiple innate immune processes in the resolution of otitis media. Innate Immun. 2015 Feb; 21(2):203-14. PMID: 24652041; PMCID: PMC4379507.
    35. Brand Y, Sung M, Chavez E, Wei E, Pak KK, Housley GD, Bodmer D, Ryan AF. Neural cell adhesion molecule L1 modulates type I but not type II inner ear spiral ganglion neurite outgrowth in an in vitro alternate choice assay. J Mol Neurosci. 2013 Nov; 51(3):663-70. PMID: 23760987; PMCID: PMC3825176.
    36. Kurabi A, Pak K, Dang X, Coimbra R, Eliceiri BP, Ryan AF, Baird A. Ecrg4 attenuates the inflammatory proliferative response of mucosal epithelial cells to infection. PLoS One. 2013; 8(4):e61394. PMID: 23626679; PMCID: PMC3634077.
    37. Housley GD, Morton-Jones R, Vlajkovic SM, Telang RS, Paramananthasivam V, Tadros SF, Wong AC, Froud KE, Cederholm JM, Sivakumaran Y, Snguanwongchai P, Khakh BS, Cockayne DA, Thorne PR, Ryan AF. ATP-gated ion channels mediate adaptation to elevated sound levels. Proc Natl Acad Sci U S A. 2013 Apr 30; 110(18):7494-9. PMID: 23592720; PMCID: PMC3645545.
    38. Li JD, Hermansson A, Ryan AF, Bakaletz LO, Brown SD, Cheeseman MT, Juhn SK, Jung TT, Lim DJ, Lim JH, Lin J, Moon SK, Post JC. Panel 4: Recent advances in otitis media in molecular biology, biochemistry, genetics, and animal models. Otolaryngol Head Neck Surg. 2013 Apr; 148(4 Suppl):E52-63. PMID: 23536532; PMCID: PMC4006668.
    39. Kondo K, Pak K, Chavez E, Mullen L, Euteneuer S, Ryan AF. Changes in responsiveness of rat spiral ganglion neurons to neurotrophins across age: differential regulation of survival and neuritogenesis. Int J Neurosci. 2013 Jul; 123(7):465-75. PMID: 23301942; PMCID: PMC3897193.
    40. Yan D, Zhu Y, Walsh T, Xie D, Yuan H, Sirmaci A, Fujikawa T, Wong AC, Loh TL, Du L, Grati M, Vlajkovic SM, Blanton S, Ryan AF, Chen ZY, Thorne PR, Kachar B, Tekin M, Zhao HB, Housley GD, King MC, Liu XZ. Mutation of the ATP-gated P2X(2) receptor leads to progressive hearing loss and increased susceptibility to noise. Proc Natl Acad Sci U S A. 2013 Feb 05; 110(6):2228-33. PMID: 23345450.
      View in: PubMed
    41. Xie J, Pak K, Evans A, Kamgar-Parsi A, Fausti S, Mullen L, Ryan AF. Neurotrophins differentially stimulate the growth of cochlear neurites on collagen surfaces and in gels. Neural Regen Res. 2013; 8(17):1541-1550. PMID: 24459465.
      View in: PubMed
    42. Euteneuer S, Yang KH, Chavez E, Leichtle A, Loers G, Olshansky A, Pak K, Schachner M, Ryan AF. Glial cell line-derived neurotrophic factor (GDNF) induces neuritogenesis in the cochlear spiral ganglion via neural cell adhesion molecule (NCAM). Mol Cell Neurosci. 2013 May; 54:30-43. PMID: 23262364; PMCID: PMC3857543.
    43. Jin Y, Kondo K, Ushio M, Kaga K, Ryan AF, Yamasoba T. Developmental changes in the responsiveness of rat spiral ganglion neurons to neurotrophic factors in dissociated culture: differential responses for survival, neuritogenesis and neuronal morphology. Cell Tissue Res. 2013 Jan; 351(1):15-27. PMID: 23149719; PMCID: PMC3577061.
    44. Masuda M, Pak K, Chavez E, Ryan AF. TFE2 and GATA3 enhance induction of POU4F3 and myosin VIIa positive cells in nonsensory cochlear epithelium by ATOH1. Dev Biol. 2012 Dec 01; 372(1):68-80. PMID: 22985730; PMCID: PMC3483650.
    45. Cederholm JM, Froud KE, Wong AC, Ko M, Ryan AF, Housley GD. Differential actions of isoflurane and ketamine-based anaesthetics on cochlear function in the mouse. Hear Res. 2012 Oct; 292(1-2):71-9. PMID: 22960466; PMCID: PMC3582347.
    46. Souter M, Eastwood H, Marovic P, Kel G, Wongprasartsuk S, Ryan AF, O'Leary SJ. Systemic immunity influences hearing preservation in cochlear implantation. Otol Neurotol. 2012 Jun; 33(4):532-8. PMID: 22470051; PMCID: PMC3897157.
    47. Husseman J, Palacios SD, Rivkin AZ, Oehl H, Ryan AF. The role of vascular endothelial growth factors and fibroblast growth factors in angiogenesis during otitis media. Audiol Neurootol. 2012; 17(3):148-54. PMID: 22104377; PMCID: PMC3238030.
    48. Brand Y, Setz C, Levano S, Listyo A, Chavez E, Pak K, Sung M, Radojevic V, Ryan AF, Bodmer D. Simvastatin protects auditory hair cells from gentamicin-induced toxicity and activates Akt signaling in vitro. BMC Neurosci. 2011 Nov 14; 12:114. PMID: 22082490; PMCID: PMC3250952.
    49. Mullen LM, Pak KK, Chavez E, Kondo K, Brand Y, Ryan AF. Ras/p38 and PI3K/Akt but not Mek/Erk signaling mediate BDNF-induced neurite formation on neonatal cochlear spiral ganglion explants. Brain Res. 2012 Jan 09; 1430:25-34. PMID: 22119396; PMCID: PMC3242932.
    50. Barclay M, Ryan AF, Housley GD. Type I vs type II spiral ganglion neurons exhibit differential survival and neuritogenesis during cochlear development. Neural Dev. 2011 Oct 11; 6:33. PMID: 21989106; PMCID: PMC3207869.
    51. Ahmad ZK, Brown CM, Cueva RA, Ryan AF, Doherty JK. ErbB expression, activation, and inhibition with lapatinib and tyrphostin (AG825) in human vestibular schwannomas. Otol Neurotol. 2011 Jul; 32(5):841-7. PMID: 21659924; PMCID: PMC3584631.
    52. Leichtle A, Lai Y, Wollenberg B, Wasserman SI, Ryan AF. Innate signaling in otitis media: pathogenesis and recovery. Curr Allergy Asthma Rep. 2011 Feb; 11(1):78-84. PMID: 21049294; PMCID: PMC3020300.
    53. Ebmeyer J, Leichtle A, Hernandez M, Ebmeyer U, Husseman J, Pak K, Sudhoff H, Broide D, Wasserman SI, Ryan AF. TNFA deletion alters apoptosis as well as caspase 3 and 4 expression during otitis media. BMC Immunol. 2011 Jan 26; 12:12. PMID: 21269505; PMCID: PMC3040143.
    54. Leichtle A, Hernandez M, Lee J, Pak K, Webster NJ, Wollenberg B, Wasserman SI, Ryan AF. The role of DNA sensing and innate immune receptor TLR9 in otitis media. Innate Immun. 2012 Feb; 18(1):3-13. PMID: 21239460; PMCID: PMC4041324.
    55. Brown CM, Ahmad ZK, Ryan AF, Doherty JK. Estrogen receptor expression in sporadic vestibular schwannomas. Otol Neurotol. 2011 Jan; 32(1):158-62. PMID: 21099731; PMCID: PMC3073320.
    56. Lai Y, Cogen AL, Radek KA, Park HJ, Macleod DT, Leichtle A, Ryan AF, Di Nardo A, Gallo RL. Activation of TLR2 by a small molecule produced by Staphylococcus epidermidis increases antimicrobial defense against bacterial skin infections. J Invest Dermatol. 2010 Sep; 130(9):2211-21. PMID: 20463690; PMCID: PMC2922455.
    57. Ahmad Z, Brown CM, Patel AK, Ryan AF, Ongkeko R, Doherty JK. Merlin knockdown in human Schwann cells: clues to vestibular schwannoma tumorigenesis. Otol Neurotol. 2010 Apr; 31(3):460-6. PMID: 20195187; PMCID: PMC2873969.
    58. Leichtle A, Hernandez M, Ebmeyer J, Yamasaki K, Lai Y, Radek K, Choung YH, Euteneuer S, Pak K, Gallo R, Wasserman SI, Ryan AF. CC chemokine ligand 3 overcomes the bacteriocidal and phagocytic defect of macrophages and hastens recovery from experimental otitis media in TNF-/- mice. J Immunol. 2010 Mar 15; 184(6):3087-97. PMID: 20164426; PMCID: PMC2929161.
    59. Barclay M, Julien JP, Ryan AF, Housley GD. Type III intermediate filament peripherin inhibits neuritogenesis in type II spiral ganglion neurons in vitro. Neurosci Lett. 2010 Jul 05; 478(2):51-5. PMID: 20132868; PMCID: PMC2902986.
    60. Ebmeyer J, Ebmeyer U, Pak K, Sudhoff H, Broide D, Ryan AF, Wasserman S. Reconstitution of the mast cell population in W/Wv mice. Otol Neurotol. 2010 Jan; 31(1):42-7. PMID: 19752767; PMCID: PMC2796298.
    61. Lai Y, Di Nardo A, Nakatsuji T, Leichtle A, Yang Y, Cogen AL, Wu ZR, Hooper LV, Schmidt RR, von Aulock S, Radek KA, Huang CM, Ryan AF, Gallo RL. Commensal bacteria regulate Toll-like receptor 3-dependent inflammation after skin injury. Nat Med. 2009 Dec; 15(12):1377-82. PMID: 19966777; PMCID: PMC2880863.
    62. Leichtle A, Hernandez M, Pak K, Webster NJ, Wasserman SI, Ryan AF. The toll-Like receptor adaptor TRIF contributes to otitis media pathogenesis and recovery. BMC Immunol. 2009 Aug 05; 10:45. PMID: 19656404; PMCID: PMC2736931.
    63. Leichtle A, Hernandez M, Pak K, Yamasaki K, Cheng CF, Webster NJ, Ryan AF, Wasserman SI. TLR4-mediated induction of TLR2 signaling is critical in the pathogenesis and resolution of otitis media. Innate Immun. 2009 Aug; 15(4):205-15. PMID: 19586996; PMCID: PMC3565845.
    64. Ryan AF, Mullen LM, Doherty JK. Cellular targeting for cochlear gene therapy. Adv Otorhinolaryngol. 2009; 66:99-115. PMID: 19494575; PMCID: PMC4379510.
    65. Ryan AF, Dazert S. Gene therapy for the inner ear: challenges and promises. Adv Otorhinolaryngol. 2009; 66:1-12. PMID: 19494569; PMCID: PMC4332585.
    66. Hernandez M, Leichtle A, Pak K, Ebmeyer J, Euteneuer S, Obonyo M, Guiney DG, Webster NJ, Broide DH, Ryan AF, Wasserman SI. Myeloid differentiation primary response gene 88 is required for the resolution of otitis media. J Infect Dis. 2008 Dec 15; 198(12):1862-9. PMID: 18986247; PMCID: PMC3584630.
    67. Du X, Schwander M, Moresco EM, Viviani P, Haller C, Hildebrand MS, Pak K, Tarantino L, Roberts A, Richardson H, Koob G, Najmabadi H, Ryan AF, Smith RJ, Müller U, Beutler B. A catechol-O-methyltransferase that is essential for auditory function in mice and humans. Proc Natl Acad Sci U S A. 2008 Sep 23; 105(38):14609-14. PMID: 18794526; PMCID: PMC2567147.
    68. Patel AK, Alexander TH, Andalibi A, Ryan AF, Doherty JK. Vestibular schwannoma quantitative polymerase chain reaction expression of estrogen and progesterone receptors. Laryngoscope. 2008 Aug; 118(8):1458-63. PMID: 18670322; PMCID: PMC3570025.
    69. Brors D, Hansen S, Mlynski R, Volkenstein S, Aletsee C, Sendtner M, Ryan AF, Dazert S. Spiral ganglion outgrowth and hearing development in p75-deficient mice. Audiol Neurootol. 2008; 13(6):388-95. PMID: 18663291.
      View in: PubMed
    70. Doherty JK, Ongkeko W, Crawley B, Andalibi A, Ryan AF. ErbB and Nrg: potential molecular targets for vestibular schwannoma pharmacotherapy. Otol Neurotol. 2008 Jan; 29(1):50-7. PMID: 18199957.
      View in: PubMed
    71. Evans AR, Euteneuer S, Chavez E, Mullen LM, Hui EE, Bhatia SN, Ryan AF. Laminin and fibronectin modulate inner ear spiral ganglion neurite outgrowth in an in vitro alternate choice assay. Dev Neurobiol. 2007 Nov; 67(13):1721-30. PMID: 17600813.
      View in: PubMed
    72. Barclay M, Noakes PG, Ryan AF, Julien JP, Housley GD. Neuronal expression of peripherin, a type III intermediate filament protein, in the mouse hindbrain. Histochem Cell Biol. 2007 Dec; 128(6):541-50. PMID: 17899157.
      View in: PubMed
    73. Dazert S, Schick B, Hartensuer R, Volkenstein S, Aletsee C, Hansen S, Shehata-Dieler WE, Eigenthaler M, Walter U, Ryan AF, Brors D. Hearing development and spiral ganglion neurite growth in VASP deficient mice. Brain Res. 2007 Oct 31; 1178:73-82. PMID: 17920567.
      View in: PubMed
    74. Hedlund M, Tangvoranuntakul P, Takematsu H, Long JM, Housley GD, Kozutsumi Y, Suzuki A, Wynshaw-Boris A, Ryan AF, Gallo RL, Varki N, Varki A. N-glycolylneuraminic acid deficiency in mice: implications for human biology and evolution. Mol Cell Biol. 2007 Jun; 27(12):4340-6. PMID: 17420276; PMCID: PMC1900035.
    75. Greenwood D, Jagger DJ, Huang LC, Hoya N, Thorne PR, Wildman SS, King BF, Pak K, Ryan AF, Housley GD. P2X receptor signaling inhibits BDNF-mediated spiral ganglion neuron development in the neonatal rat cochlea. Development. 2007 Apr; 134(7):1407-17. PMID: 17329369.
      View in: PubMed
    76. Schwander M, Sczaniecka A, Grillet N, Bailey JS, Avenarius M, Najmabadi H, Steffy BM, Federe GC, Lagler EA, Banan R, Hice R, Grabowski-Boase L, Keithley EM, Ryan AF, Housley GD, Wiltshire T, Smith RJ, Tarantino LM, Müller U. A forward genetics screen in mice identifies recessive deafness traits and reveals that pejvakin is essential for outer hair cell function. J Neurosci. 2007 Feb 28; 27(9):2163-75. PMID: 17329413.
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    77. Furukawa M, Ebmeyer J, Pak K, Austin DA, Melhus A, Webster NJ, Ryan AF. Jun N-terminal protein kinase enhances middle ear mucosal proliferation during bacterial otitis media. Infect Immun. 2007 May; 75(5):2562-71. PMID: 17325051; PMCID: PMC1865762.
    78. Chung WH, Pak K, Lin B, Webster N, Ryan AF. A PI3K pathway mediates hair cell survival and opposes gentamicin toxicity in neonatal rat organ of Corti. J Assoc Res Otolaryngol. 2006 Dec; 7(4):373-82. PMID: 17053865; PMCID: PMC2504631.
    79. Ryan AF, Ebmeyer J, Furukawa M, Pak K, Melhus A, Wasserman SI, Chung WH. Mouse models of induced otitis media. Brain Res. 2006 May 26; 1091(1):3-8. PMID: 16764829.
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    80. Ryan AF, Wittig J, Evans A, Dazert S, Mullen L. Environmental micro-patterning for the study of spiral ganglion neurite guidance. Audiol Neurootol. 2006; 11(2):134-43. PMID: 16439836.
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    81. Huang LC, Ryan AF, Cockayne DA, Housley GD. Developmentally regulated expression of the P2X3 receptor in the mouse cochlea. Histochem Cell Biol. 2006 Jun; 125(6):681-92. PMID: 16341871.
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    82. Ebmeyer J, Furukawa M, Pak K, Ebmeyer U, Sudhoff H, Broide D, Ryan AF, Wasserman S. Role of mast cells in otitis media. J Allergy Clin Immunol. 2005 Nov; 116(5):1129-35. PMID: 16275387.
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    83. Rivkin AZ, Palacios SD, Pak K, Bennett T, Ryan AF. The role of Fas-mediated apoptosis in otitis media: observations in the lpr/lpr mouse. Hear Res. 2005 Sep; 207(1-2):110-6. PMID: 15978756.
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    84. Ryan AF, Jung TT, Juhn SK, Li JD, Andalibi A, Lin J, Bakaletz LO, Post CJ, Ehrlich GD. Recent advances in otitis media. 4A. Molecular biology. Ann Otol Rhinol Laryngol Suppl. 2005 Jan; 194:42-9. PMID: 15700934.
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    85. Ryan AF, Jung TT, Juhn SK, Li JD, Andalibi A, Lin J, Bakaletz LO, Post CJ, Ehrlich GD. Recent advances in otitis media. 4C. Interaction between middle ear and inner ear in otitis media. Ann Otol Rhinol Laryngol Suppl. 2005 Jan; 194:56-9. PMID: 15700936.
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    86. Ryan AF, Jung TT, Juhn SK, Li JD, Andalibi A, Lin J, Bakaletz LO, Post CJ, Ehrlich GD. Recent advances in otitis media. 4B. Biochemistry. Ann Otol Rhinol Laryngol Suppl. 2005 Jan; 194:50-5. PMID: 15700935.
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    87. Wittig JH, Ryan AF, Asbeck PM. A reusable microfluidic plate with alternate-choice architecture for assessing growth preference in tissue culture. J Neurosci Methods. 2005 May 15; 144(1):79-89. PMID: 15848242.
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    88. Palacios SD, Pak K, Rivkin AZ, Kayali AG, Austen D, Aletsee C, Melhus A, Webster NJ, Ryan AF. Role of p38 mitogen-activated protein kinase in middle ear mucosa hyperplasia during bacterial otitis media. Infect Immun. 2004 Aug; 72(8):4662-7. PMID: 15271927; PMCID: PMC470656.
    89. Melhus A, Ryan AF. Effects of amoxicillin on cytokine and osteocalcin expression in bone tissue during experimental acute otitis media. Cytokine. 2004 Mar 21; 25(6):254-9. PMID: 15036240.
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    90. Barash IA, Mathew L, Ryan AF, Chen J, Lieber RL. Rapid muscle-specific gene expression changes after a single bout of eccentric contractions in the mouse. Am J Physiol Cell Physiol. 2004 Feb; 286(2):C355-64. PMID: 14561590.
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    91. Melhus A, Ryan AF. A mouse model for acute otitis media. APMIS. 2003 Oct; 111(10):989-94. PMID: 14616553.
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    92. Bodmer D, Brors D, Bodmer M, Pak K, Ryan AF. Fas ligand expression in the organ of Corti. Audiol Neurootol. 2003 Sep-Oct; 8(5):243-9. PMID: 12904679.
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    93. Brors D, Bodmer D, Pak K, Aletsee C, Schäfers M, Dazert S, Ryan AF. EphA4 provides repulsive signals to developing cochlear ganglion neurites mediated through ephrin-B2 and -B3. J Comp Neurol. 2003 Jul 14; 462(1):90-100. PMID: 12761826.
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    94. Wang JC, Raybould NP, Luo L, Ryan AF, Cannell MB, Thorne PR, Housley GD. Noise induces up-regulation of P2X2 receptor subunit of ATP-gated ion channels in the rat cochlea. Neuroreport. 2003 May 06; 14(6):817-23. PMID: 12858039.
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    95. Aletsee C, Brors D, Mlynski R, Ryan AF, Dazert S. Branching of spiral ganglion neurites is induced by focal application of fibroblast growth factor-1. Laryngoscope. 2003 May; 113(5):791-6. PMID: 12792312.
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    96. Bodmer D, Brors D, Pak K, Bodmer M, Ryan AF. Gentamicin-induced hair cell death is not dependent on the apoptosis receptor Fas. Laryngoscope. 2003 Mar; 113(3):452-5. PMID: 12616196.
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    97. Skinner LJ, Enée V, Beurg M, Jung HH, Ryan AF, Hafidi A, Aran JM, Dulon D. Contribution of BK Ca2+-activated K+ channels to auditory neurotransmission in the Guinea pig cochlea. J Neurophysiol. 2003 Jul; 90(1):320-32. PMID: 12611976.
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    98. Brors D, Aletsee C, Dazert S, Huverstuhl J, Ryan AF, Bodmer D. Clostridium difficile toxin B, an inhibitor of the small GTPases Rho, Rac and Cdc42, influences spiral ganglion neurite outgrowth. Acta Otolaryngol. 2003 Jan; 123(1):20-5. PMID: 12625568.
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    99. Ryan AF. The cell cycle and the development and regeneration of hair cells. Curr Top Dev Biol. 2003; 57:449-66. PMID: 14674489.
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    100. Palacios SD, Pak K, Kayali AG, Rivkin AZ, Aletsee C, Melhus A, Webster NJ, Ryan AF. Participation of Ras and extracellular regulated kinase in the hyperplastic response of middle-ear mucosa during bacterial otitis media. J Infect Dis. 2002 Dec 15; 186(12):1761-9. PMID: 12447762.
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    101. Bodmer D, Gloddek B, Ryan AF, Huverstuhl J, Brors D. Inhibition of the c-Jun N-terminal kinase signaling pathway influences neurite outgrowth of spiral ganglion neurons in vitro. Laryngoscope. 2002 Nov; 112(11):2057-61. PMID: 12439181.
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    102. Gloddek B, Bodmer D, Brors D, Keithley EM, Ryan AF. Induction of MHC class II antigens on cells of the inner ear. Audiol Neurootol. 2002 Nov-Dec; 7(6):317-23. PMID: 12463193.
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    103. Bodmer D, Brors D, Pak K, Gloddek B, Ryan A. Rescue of auditory hair cells from aminoglycoside toxicity by Clostridium difficile toxin B, an inhibitor of the small GTPases Rho/Rac/Cdc42. Hear Res. 2002 Oct; 172(1-2):81-6. PMID: 12361869.
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    104. Bodmer D, Brors D, Pak K, Keithley EM, Mullen L, Ryan AF, Gloddek B. Inflammatory signals increase Fas ligand expression by inner ear cells. J Neuroimmunol. 2002 Aug; 129(1-2):10-7. PMID: 12161015.
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    105. Brors D, Aletsee C, Schwager K, Mlynski R, Hansen S, Schäfers M, Ryan AF, Dazert S. Interaction of spiral ganglion neuron processes with alloplastic materials in vitro(1). Hear Res. 2002 May; 167(1-2):110-21. PMID: 12117535.
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    106. Ryan AF. Molecular studies of hair cell development and survival. Audiol Neurootol. 2002 May-Jun; 7(3):138-40. PMID: 12053133.
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    107. Palacios SD, Pak K, Rivkin AZ, Bennett T, Ryan AF. Growth factors and their receptors in the middle ear mucosa during otitis media. Laryngoscope. 2002 Mar; 112(3):420-3. PMID: 12148846.
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    108. Aletsee C, Brors D, Palacios S, Pak K, Mullen L, Dazert S, Ryan AF. The effects of laminin-1 on spiral ganglion neurons are dependent on the MEK/ERK signaling pathway and are partially independent of Ras. Hear Res. 2002 Feb; 164(1-2):1-11. PMID: 11950519.
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    109. Ryan AF, Harris JP, Keithley EM. Immune-mediated hearing loss: basic mechanisms and options for therapy. Acta Otolaryngol Suppl. 2002; (548):38-43. PMID: 12211356.
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    110. Ryan AF, Pak K, Low W, Battaglia A, Mullen L, Harris JP, Keithley EM. Immunological damage to the inner ear: current and future therapeutic strategies. Adv Otorhinolaryngol. 2002; 59:66-74. PMID: 11885663.
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    111. Ryan AF, Bennett T. Nitric oxide contributes to control of effusion in experimental otitis media. Laryngoscope. 2001 Feb; 111(2):301-5. PMID: 11210879.
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    112. Ryan AF. Summary: progress in inner-ear immunology. Ann N Y Acad Sci. 1997 Dec 29; 830:326-9. PMID: 9616693.
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    113. Ryan AF, Gloddek B, Harris JP. Lymphocyte trafficking to the inner ear. Ann N Y Acad Sci. 1997 Dec 29; 830:236-42. PMID: 9616682.
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