Emilio Flano, Ph.D. :: Nationwide Children's Hospital, Columbus, Ohio

Emilio Flano, Ph.D.

Emilio  Flano, Ph.D.

Center for Vaccines and Immunity
Principal Investigator

Contact Information

The Research Institute at Nationwide Children's Hospital
700 Childrens Drive WA4015
Columbus, Ohio 43205 [ map ]
PH: (614) 722.2735
FX: (614) 722.3680
E-mail Me

Biography

Emilio Flano, PhD, is a Principal Investigator in the Center for Vaccines and Immunity and the Director of the Flow Cytometry Core Laboratory at The Research Institute at Nationwide Children's Hospital. He is an Associate Professor of Pediatrics at the The Ohio State University College of Medicine. His research program focuses on the mechanisms of immune protection against viral infections, with an emphasis on respiratory viruses.

View CV »

Gender:

  • Male

Languages Spoken:

  • English

Research Interests

Research Center:

Areas of Interest:

  • Infectious diseases are a major health priority. My work aims to understand the mechanisms of immune protection against viral infections, which a focus on infections initiated in the respiratory tract. To accomplish this, we use a several models of viral infection including murine ?-herpesvirus 68, influenza, respiratory syncytial virus and Newcastle disease virus. Two are our main research interests.

    First, to understand how T cells are recruited into the respiratory tract in response to infection. The lung is the major portal of entry for microorganisms in the body. However, we still do not know how T cells are specifically recruited to the lung and into the airways during infection. We propose that type I interferon signaling mediates T cell trafficking into the airways following virus infection of the respiratory epithelium.

    Second, another goal in our laboratory is to understand how protective T cell memory can be elicited and maintained long-term. To develop safe and effective vaccines we need a thorough knowledge of the mechanisms that regulate immunological memory. Although long-lived, protective T cell memory wanes with time after viral clearance. We also know that high-level antigen persistence is detrimental for T cell memory and function. More than 95% of the human adult population shows evidence of life-long latent persistent infection by herpesviruses. However, only a minority of these individuals develop herpesvirus-associated diseases. We propose that understanding the cellular and molecular mechanisms that maintain life-long protective T cell responses during herpesvirus persistence will be instrumental for elucidating how long-lasting cellular immunity is generated and maintained.
  • Infectious diseases are a major health priority. My work aims to understand the mechanisms of immune protection against viral infections, which a focus on infections initiated in the respiratory tract. To accomplish this, we use a several models of viral infection including murine ?-herpesvirus 68, influenza, respiratory syncytial virus and Newcastle disease virus.

Education and Training

Post Doctoral

  • University of León
    Date Completed: 06/30/1996

Post Doctoral

  • St. Jude Children’s Research Hospital - Postdoctoral Research Associate
    Date Completed: 06/30/2000

Professional Experience

2010–present

  • Associate Professor, Department of Pediatrics, The Ohio State University School of Medicine, Columbus, OH.

2007–present

  • Director, Flow Cytometry Core Facility, The Research Institute at Nationwide Children’s Hospital, Columbus, OH.

2004–present

  • Principal Investigator, Center for Vaccines and Immunity, The Research Institute at Nationwide Children’s Hospital, Columbus, OH.

2004–2010

  • Assistant Professor, Department of Pediatrics, The Ohio State University School of Medicine, Columbus, OH.

2001–2004

  • Research Assistant Member, Trudeau Institute, Saranac Lake, NY.

2000–2001

  • Research Associate, Trudeau Institute, Saranac Lake, NY. 

1997–2000

  • Postdoctoral Research Associate, Department of Immunology, St. Jude Children’s Research Hospital, Memphis, TN.

Publications

  • Cao RG, Suarez NM, Obermoser G, Lopez SM, Flano E, Mertz SE, Albrecht RA, Garcia-Sastre A, Mejias A, Xu H, Qin H, Blankenship D, Palucka K, Pascual V, Ramilo O. 2014. Differences in antibody responses between TIV and LAIV influenza vaccines correlate with the kinetics and magnitude of interferon signaling in children.  J Infect Dis. Vol. Epub ahead of print Feb 4, no. February: ena.
  • McNally B, Ye F, Willette M, Flanoe E. 2013. Local blockade of epithelial PDL-1 in the airways enhances T cell function and viral clearance during influenza virus infection.  J Virol. Vol. 87, no. 23. (December): 12916-24.
  • Betebibel, S.-E., Lopez, S., Obermoser, G., Schmitt, N., Mueller, C., Harrod, C., Flano, E., Mejias, A., Albrecht, R.A., Blankenship, D., Xu, H., Pascual, V, Bancherau, J., Garcia-Sastre, A., Palucka, K., Ramilo, O., Ueno, H. 2013. Inductin of ICOS+CXCR3+CXCR5+ TH cells correlates with antibody responses to influenza vaccination.  Sci Transl Med. Vol. 5, no. 176. (March): e176ra32.
  • Ioannidis I, Ye F, McNally B, Willette M, Flaño E. 2013. TLR expression and induction of type I and type III interferons in primary airway epithelial cells.  J Virol. Vol. 87, no. 6. (January): 3261-3270.
  • Cyktor, J, Caruthers, B, Stromberg, P, Flano, E, Pircher, H, Turner, E. 2013. KLRG1 deficiency significantly enhances survival after Mycobacterium tuberculosis infection.  Infec Immun. Vol. 81, no. 4. (January): 1090-9.
  • McNally, B., Willette, M., Partida-Sanchez, S., Flaño, E. 2012. Intranasal administration of dsRNA induces interferon-? receptor-dependent accumulation of antigen experienced T cells in the airways.  Plos One. Vol. 7, no. 12. (December): ee51351.
  • Fang, Y. Turner, J., Flaño E. 2012. 2012. Contribution of Pulmonary KLRG1 high and KLRG1 low CD8 T Cells to Effector and Memory Responses during Infuenza Virus Infection.  Journal of Immunology. Vol. 189, no. 11. (October): 5206-11.
  • Aeffner F, Bratasz A, Flaño E, Powell KA, Davis IC. 2012. Post-Infection A77-1726 Treatment Improves Cardiopulmonary Function in H1N1 Influenza-Infected Mice.  Am J Respir Cell Mol Biol.. Vol. 47, no. 4. (June): 543-51.
  • Ioannidis I, McNally B, Willette M, Peeples ME, Chaussabel D, Durbin JE, Ramilo O, Mejias A, Flano E. 2012. Plasticity and virus-specificity of the airway epithelial cell immune response during respiratory viral infection.  J Virol. Vol. 86, no. 10. (March): 5422-36.
  • Freeman,Michael,L; Burkum,Claire,E; Lanzer,Kathleen,G; Jensen,Meghan,K; Ahmed,Mushtaq; Yager,Eric,J; Flano,Emilio; Winslow,Gary,M; Woodland,David,L; Blackman,Marcia,A. 2011. Cutting Edge: Activation of Virus-Specific CD4 T Cells throughout gamma-Herpesvirus Latency.  JOURNAL OF IMMUNOLOGY. Vol. 187, no. 12. (December): 6180-6184.
  • Freeman ML, Burkum CE, Lanzer KG, Jensen MK, Ahmed M, Yager EJ, Flano E, Winslow GM, Woodland DL, Blackman MA. 2011. Cutting Edge: Activation of virus-specific CD4 T cells throughout gamma-herpesvirus latency.  J Immunol. Vol. 187, no. Nov 11 ePub ahead of print. (November): 6180-6184.
  • Cush,Stephanie,S; Flano,Emilio. 2011. KLRG1(+)NKG2A(+) CD8 T Cells Mediate Protection and Participate in Memory Responses during gamma-Herpesvirus Infection.  JOURNAL OF IMMUNOLOGY. Vol. 186, no. 7. (April): 4051-4058.
  • Jewell,Nancy,A; Cline,Troy; Mertz,Sara,E; Smirnov,Sergey,V; Flano,Emilio; Schindler,Christian; Grieves,Jessica,L; Durbin,Russell,K; Kotenko,Sergei,V; Durbin,Joan,E. 2011. Lambda Interferon Is the Predominant Interferon Induced by Influenza A Virus Infection In Vivo (vol 84, pg 11515, 2010).  JOURNAL OF VIROLOGY. Vol. 85, no. 1. (January): 650-650.
  • Cush S.S. and Flaño E. 2011. KLRG1+NKG2A+ CD8 T cells mediate protection and participate in memory responses during ?-Herpesvirus infection.  J Immunol. Vol. 7, no. 186. (January): 4051-4058.
  • Weslow-Schmidt,Janet; Ye,Fang; Cush,Stephanie,S; Stuller,Kathleen,A; Blackman,Marcia,A; Flano,Emilio. 2010. Dendritic Cells Loaded with Tumor B Cells Elicit Broad Immunity against Murine Gammaherpesvirus 68 but Fail To Prevent Long-Term Latency.  JOURNAL OF VIROLOGY. Vol. 84, no. 17. (September): 8975-8979.
  • Stuller,K,A; CUSH,S,S; Flano,E. 2010. Persistent gamma-herpesvirus infection induces a CD4 T cell response containing functionally distinct effector populations (vol 184, pg 3850, 2010).  JOURNAL OF IMMUNOLOGY. Vol. 184, no. 12. (June): 7315-7315.
  • Stuller,Kathleen,A; Cush,Stephanie,S; Flano,Emilio. 2010. Persistent gamma-Herpesvirus Infection Induces a CD4 T Cell Response Containing Functionally Distinct Effector Populations.  JOURNAL OF IMMUNOLOGY. Vol. 184, no. 7. (April): 3850-3856.
  • Weslow-Schmidt, J., Ye, F., Cush, S.S., Stuller, K.A., Blackman, M.A., Flaño, E. 2010. Dendritic cells loaded with tumor B cells elicit broad immunity against murine gamma-herpesvirus 68 but fail to prevent long-term latency.  J Virol. Vol. 84, no. 17. (January): 8975-8979.
  • Jewell NA, Cline T, Mertz SE, Smirnov SV, Flano E, Schindler C, Grieves JL, Durbin RK, Kotenko SV, Durbin JE. 2010. Lambda interferon is the predominant interferon induced by influenza A virus infection in vivo.  J Virol. Vol. 84, no. 21. (January): 11515-11522.
  • Stuller, K.A., Cush, S.S. and Flaño, E. 2010. Persistent ?-herpesvirus infection induces a CD4 T cell response containing functionally distinct effector populations.  J Immunol. Vol. 184, no. 7. (January): 3850-3856.
  • Stuller,Kathleen,A; Flano,Emilio. 2009. CD4 T Cells Mediate Killing during Persistent Gammaherpesvirus 68 Infection.  JOURNAL OF VIROLOGY. Vol. 83, no. 9. (May): 4700-4703.
  • Cush,Stephanie,S; Flano,Emilio. 2009. Protective Antigen-Independent CD8 T Cell Memory Is Maintained during gamma-Herpesvirus Persistence.  JOURNAL OF IMMUNOLOGY. Vol. 182, no. 7. (April): 3995-4004.
  • Hwang,Seungmin; Kim,Kyeong,Seon; Flano,Emilio; Wu,Ting-Ting; Tong,Leming,M; Park,Ann,N; Song,Moon,Jung; Sanchez,David,Jesse; O'Connell,Ryan,M; Cheng,Genhong; Sun,Ren. 2009. Conserved Herpesviral Kinase Promotes Viral Persistence by Inhibiting the IRF-3-Mediated Type I Interferon Response.  CELL HOST & MICROBE. Vol. 5, no. 2. (February): 166-178.
  • Stuller, K.A. and Flaño, E. 2009. CD4 T cell-mediated killing in vivo during persistent gammaherpesvirus 68 infection.  J Virol. Vol. 83, no. January: 4700-4703.
  • Cush, S.S., Flaño, E. 2009. Protective antigen-independent CD8 T cell memory is maintained during ?-herpesvirus persistence.  J Immunol. Vol. 182, no. January: 3995-4004.
  • Hwang, S., Kim, K.S., Flano, E., Wu, T., Tong, L.M., Park, A.N., Song, M.J., Sanchez, D.J., O’Connell, R.M., Cheng, G. and Sun, R. 2009. Conserved herpesviral kinase promotes viral persistence by inhibiting the IRF-3-mediated type I interferon response.  Cell Host & Microbe. Vol. 5, no. January: 166-178.
  • Flano,Emilio. 2008. Viral Immunity: It Takes Two to Tango.  VIRAL IMMUNOLOGY. Vol. 21, no. 3. (September): 281-283.
  • Weslow-Schmidt,Janet,L; Jewell,Nancy,A; Mertz,Sara,E; Simas,J,Pedro; Durbin,Joan,E; Flano,Emilio. 2007. Type I interferon inhibition and dendritic cell activation during gammaherpesvirus respiratory infection.  JOURNAL OF VIROLOGY. Vol. 81, no. 18. (September): 9778-9789.
  • Weslow-Schmidt JL, Jewell NA, Mertz SE, Simas JP, Durbin JE, Flaño E. 2007. Type I interferon inhibition and dendritic cell activation during gammaherpesvirus respiratory infection.  J Virol. Vol. 81, no. 18. (January): 9778-9789.
  • Cush SS, Anderson KM, Ravneberg DH, Weslow-Schmidt JL, Flaño E. 2007. Memory generation and maintenance of CD8+ T cell function during viral persistence.  J Immunol. Vol. 179, no. 1. (January): 141-153.
  • Jewell NA, Vaghefi N, Mertz SE, Akter P, Peebles RS Jr, Bakaletz LO, Durbin RK, Flaño E, Durbin JE. 2007. Differential type I interferon induction by respiratory syncytial virus and influenza a virus in vivo.  J Virol. Vol. 81, no. 18. (January): 9790-9800.
  • Martinez-Sobrido, L., Gitiban, N., Fernandez-Sesma, A., Cros, J., Mertz, S.E., Jewell, N.A., Hammond, S., Flaño, E., Durbin, R.K., Garcia-Sastre, A., Durbin, J.E. 2006. Protection against Respiratory Syncytial Virus by a Recombinant Newcastle Disease Virus Vector.  J Virol. Vol. 80, no. January: 1130-1139.
  • Flaño, E., Basak, K., Woodland, D.L. and Blackman, M.A. 2005. Infection of dendritic cells by a γ-herpesvirus induces functional modulation.  J Immunol. Vol. 175, no. January: 3225-3234.
  • Flaño, E., Jia, Q., Moore, J., Woodland, D.L., Sun, R. and Blackman, M.A. 2005. Early establishment of gamma-herpesvirus latency: implications for immune control.  J Immunol. Vol. 174, no. January: 4972-4978.
  • Flaño, E., Hardy, C.L., Kim, I.J., Frankling, C., Coppola, M.A., Nguyen, P., Woodland, D.L., and Blackman, M.A. 2004. T cell reactivity during infectious mononucleosis and latent stages of γ-herpesvirus infection in mice.  J Immunol. Vol. 172, no. January: 3078-3085.
  • Kim,I,J; Flano,E; Woodland,D,L; Lund,F,E; Randall,T,D; Blackman,M,A. 2003. Maintenance of long term gamma-herpesvirus B cell latency is dependent on CD40-mediated development of memory B cells.  JOURNAL OF IMMUNOLOGY. Vol. 171, no. 2. (July): 886-892.
  • Kim, I-J., Flaño, E., Woodland, D.L., Lund, F.E., Randall, T.D. and Blackman, M.A. 2003. Maintenance of long-term ? -herpesvirus B cell latency is dependent on CD40-mediated development of memory B cells.  J Immunol. Vol. 171, no. January: 886-892.
  • Flaño, E., Kim, I-J., Woodland, D.L. and Blackman, M.A. 2003. Differential γ -herpesvirus distribution in distinct anatomical locations and cell subsets during persistent infection in mice.  J Immunol. Vol. 170, no. January: 3828-3834.
  • Macrae, A.I., Usherwood, E.J., Husain, S.M., Flaño, E., Kim, I.J., Woodland, D.L., Nash, A.A., Blackman, M.A., Sample, J.T. and Stewart, J.P. 2003. MHV-68 M2 protein is a B cell-associated antigen important for latency but not lymphocytosis.  J Virol. Vol. 77, no. January: 9700-9709.
  • Blackman,M,A; Flano,E. 2002. Persistent gamma-herpesvirus infections: What can we learn from an experimental mouse model?.  JOURNAL OF EXPERIMENTAL MEDICINE. Vol. 195, no. 7. (April): F29-F32.
  • Kim, I.-J., Flaño, E., Woodland, D.L., Blackman, M.A. 2002. Antibody-mediated control of persistent ? -herpesvirus infection.  J Immunol. Vol. 168, no. January: 3958-3964.
  • Flaño, E., Kim, I.J., Woodland, D.L., and Blackman, M.A. 2002. ? -Herpesvirus latency is preferentially maintained in splenic germinal center and memory B cells.  J Exp Med. Vol. 196, no. January: 1363-1372.
  • Blackman, M.A. and Flaño, E. 2002. Persistent gammaherpesvirus infections: what can we learn from an experimental mouse model?.  J Exp Med. Vol. 195, no. January: F29-F32.
  • Flano,E; Woodland,D,L; Blackman,M,A; Doherty,P,C. 2001. Analysis of virus-specific CD4(+) T cells during long-term gammaherpesvirus infection.  JOURNAL OF VIROLOGY. Vol. 75, no. 16. (August): 7744-7748.
  • Flaño, E., Woodland, D.L., Blackman, M.A., Doherty, P.C. 2001. Analysis of virus-specific CD4+ T cells during long-term gammaherpesvirus infection.  J Virol. Vol. 75, no. January: 7744-7748.
  • Hardy, C.L., Flaño, E., Cardin, R.C., Kim, I.-J., Nguyen, P., King, S., Woodland, D.L., Blackman, M.A. 2001. Factors controlling CD8+ T cell activation associated with murine γ-herpesvirus-induced infectious mononucleosis.  Viral Immunol. Vol. 14, no. January: 391-402.
  • Flaño, E, Woodland, D.L., Blackman, M.A. 2001. A mouse model for infectious mononucleosis.  Immunologic Research. Vol. 24, no. January: 353-369.
  • Woodland, D. L., Flaño, E., Usherwood, E. J., Liu, L., Kim, I.-J., Husain, S. M., Sample, J. T., and Blackman, M. A. 2001. Antigen expression during murine ? -herpesvirus infection.  Immunobiology. Vol. 204, no. January: 649-658.
  • Woodland, D.L., Usherwood, E.J., Liu, L., Flaño, E., Kim, I.-J., Blackman, M.A. 2001. Vaccination against murine ? -herpesvirus infection.  Viral Immunol. Vol. 14, no. January: 217-226.
  • Blackman,M,A; Flano,E; Usherwood,E; Woodland,D,L. 2000. Murine gamma-herpesvirus-68: a mouse model for infectious mononucleosis?.  MOLECULAR MEDICINE TODAY. Vol. 6, no. 12. (December): 488-490.
  • McIntosh,D; Austin,B; Flano,E; Villena,A; Martinez-Pereda,J,A; Tarazona,J,V. 2000. Lack of uptake of Renibacterium salmoninarum by gill epithelia of rainbow trout.  JOURNAL OF FISH BIOLOGY. Vol. 56, no. 5. (May): 1053-1061.
  • Flaño, E., Husain, S.M., Sample, J.T., Woodland, D.L., Blackman, M.A. 2000. Latent murine ?-herpesvirus infection is established in activated B cells, dendritic cells and macrophages.  J Immunol. Vol. 165, no. January: 1074-1081.
  • Blackman, M.A., Flaño, E., Usherwood, E., Woodland, D.L. 2000. Murine ?-herpesvirus-68, a model for infectious mononucleosis?.  Molec Med Today. Vol. 6, no. January: 488-490.
  • McIntosh, D., Austin, B., Flaño, E., Villena, A., Martinez-Pereda, J.A., Tarazona, J.V. 2000. Lack of uptake of Renibacteriaum salmoninarum by gill epithelia of rainbow trout (Onchorhynchus mykiss, Walbaum).  J Fish Biology. Vol. 56, no. January: 1053-1061.
  • Coppola, M.A., Flaño, E., Nguyen, P., Hardy, C.L., Cardin, R.D., Shastri, N., Woodland, D.L., Blackman, M.A. 1999. MHC-independent stimulation of CD8+ T cells in vivo during latent murine ?-herpesvirus infection.  J Immunol. Vol. 163, no. January: 1481-1489.
  • Flaño, E., Woodland, L.D., Blackman, M.A. 1999. Requirement for CD4+T cells in Vb4+CD8+ T cell activation associated with latent murine gammaherpesvirus infection.  J Immunol. Vol. 163, no. January: 3403-3408.
  • Liu, L., Flaño, E., Usherwood, E.J., Surman, S.L., Blackman, M.A., Woodland, D.L. 1999. Lytic cycle T cell epitopes are expressed in two distinct phases during MHV-68 infection.  J Immunol. Vol. 163, no. January: 868-874.
  • Schroder,M,B; Flano,E; PILSTROM,L; Jorgensen,T,O. 1998. Localisation of Ig heavy chain mRNA positive cells in Atlantic cod (Gadus morhua L) tissues identified by in situ hybridisation.  FISH & SHELLFISH IMMUNOLOGY. Vol. 8, no. 8. (November): 565-576.
  • Flano,E; Lopez-Fierro,P; Alvarez,F; Razquin,B; Villena,A. 1998. Splenic cultures from rainbow trout, Oncorhynchus mykiss: establishment and characterisation.  FISH & SHELLFISH IMMUNOLOGY. Vol. 8, no. 8. (November): 589-606.
  • Flaño, E., López-Fierro, P., Alvarez, F., Razquin, B., Villena, A. 1998. Splenic cultures from rainbow trout, Oncorhynchus mykiss: establishment and characterisation.  Fish Shellfish Immunol. Vol. 8, no. January: 589-606.
  • Schroder, M.B., Flaño, E., Pilstrom, L., Jorgensen, T.O. 1998. Localisation of plasma cells in Atlantic cod (Gadus morhua L.) tissues; identified by is situ hybridisation.  Fish Shellfish Immunol. Vol. 8, no. January: 565-576.
  • Hernanz Moral, C., Flaño del Castillo E., Lopez Fierro, P., Villena Cortes, A., Anguita Castillo, J., Cascon Soriano, A., Sanchez Salazar, M., Razquin Peralta, B., Naharro Carrasco, G. 1998. Molecular characterization of the Aeromonas hydrophila aroA gene and potential use of an auxothrophic aroA mutant as live attenuated vaccine.  Infect Immunity. Vol. 66, no. January: 1813-1821.
  • McIntosh,D; Flano,E; Grayson,T,H; Gilpin,M,L; Austin,B; Villena,A,J. 1997. Production of putative virulence factors by Renibacterium salmoninarum grown in cell culture.  MICROBIOLOGY-SGM. Vol. 143, no. October: 3349-3356.
  • Flano,E; LOPEZFIERRO,P; Razquin,B,E; Villena,A. 1997. In vitro proliferation of eosinophilic granular cells in gill cultures from rainbow trout.  FISH & SHELLFISH IMMUNOLOGY. Vol. 7, no. 7. (October): 519-521.
  • McIntosh, D., Flaño, E., Grayson, T.H., Gilpin, M.L., Austin, B., Villena, A. 1997. Production of putative virulence factors by Renibacterium salmoninarum grown in cell culture.  Microbiol. Vol. 134, no. January: 3349-3356.
  • Flaño, E., López-Fierro, P., Razquin, B., Villena, A. 1997. In vitro proliferation of eosinophilic granular cells in gill cultures from rainbow trout.  Fish Shellfish Immunol. Vol. 7, no. January: 519-521.
  • Flaño, E., López-Fierro, P., Razquin, B., Kaattari, S.L., Villena, A. 1996. Histopathology of the renal and splenic haemopoietic tissues of coho salmon, Oncorhynchus kisutch, experimentally infected with Renibacterium salmoninarum.  Diseases Aquatic Organisms. Vol. 24, no. January: 107-115.
  • Flaño, E., López-Fierro, P., Razquin, B.E., Villena, A. 1996. In vitro differentiation of eosinophilic granular cells in Renibacterium salmoninarum-infected gill cultures from rainbow trout.  Fish Shellfish Immunol. Vol. 6, no. January: 173-184.
  • Flaño, E., Alvarez, F., López-Fierro, P., Razquin, B., Villena, A., Zapata, A. 1996. In vitro and in situ characterisation of fish thymic nurse cells.  Dev Immunol. Vol. 26, no. January: 11-18.
  • Alvarez, F., Flaño, E., Castillo, A., López-Fierro, P., Razquin, B., Villena, A. 1996. Tissue distribution of barrier cells in the hematopoietic and lymphoid organs of salmonids.  Anatomical Record. Vol. 245, no. January: 17-24.
  • Flano,E; Alvarez,F; LOPEZFIERRO,P; Razquin,B,E; Villena,A,J; Zapata,A,G. 1996. In vitro and in situ characterization of fish thymic nurse cells.  DEVELOPMENTAL IMMUNOLOGY. Vol. 5, no. 1. (January): 17-24.
  • Flaño, E., Kaattari, S.L., Razquin, B., Villena, A. 1996. Histopathology of the thymus of coho salmon, Oncorhynchus kisutch, experimentally infected with Renibacterium salmoninarum.  Diseases Aquatic Organisms. Vol. 26, no. January: 11-18.
  • Alvarez,F; Flano,E; Villena,A,J; Zapata,A; Razquin,B,E. 1994. SEASONAL INTRATHYMIC ERYTHROPOIETIC ACTIVITY IN TROUT.  DEVELOPMENTAL AND COMPARATIVE IMMUNOLOGY. Vol. 18, no. 5. (September): 409-420.
  • Alvarez, F., Flaño, E., Villena, A., Zapata, A., Razquin, A. 1994. Seasonal intrathymic erythropoietic activity in trout.  Develop Comparative Immunol. Vol. 18, no. January: 409-420.
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