image001image002The Cytokine and Phagocyte Biology Lab



*                         Who and where are we?

*                         Teaching

*                         Research interest

*                         Personnel

*                         Publications

*                         Research grants

*                         Positions available:

Who and where are we?

Academic staff/Senior Scientists:

Prof. Victoriano Mulero

Prof. Alfonsa Garcia-Ayala

Prof. Jose Meseguer

Dr. Maria P. Sepulcre


Department of Cell Biology and Histology. Faculty of Biology.

University of Murcia. 30100 Murcia. SPAIN.

Contact us:

V. Mulero:, +34 868 887581

A. Garcia-Ayala:, +34 868 884968

J. Meseguer:, +34 868 884965

M.P. Sepulcre:, +34 868 883938

FAX: +34 868 883963

Go to index


BSc courses:

Biology degree
Cell Biology
Biotechnology degree
Cell Biology
Basic Cell Biology Techniques
Advanced Cell Biology Techniques
Tissue engineering 
Evolutionary immunology
Animal cell culture
Biotechnological applications of antibodies 

MSc and PhD courses:

Techniques and research in Biomedicine (Quality Mention)
 Cell Biology Techniques (1st year)
 Gene transcription and Acquisition of Cell Function (1st year)
 Phylogeny of Cytokines (Research line)
Fisheries and aquaculture
 Fish Immunity (1st year)
 The Immune Response of Fish (Research line)
Molecular Biology and Biotechnology (Quality Mention)
 Localization of Biomolecules (1st year)
 Evolution of Immunity in Vertebrates (Research line)

Go to index

Research interest

Phylogeny of immunity:

 Cytokines: functional characterisation of IL-1b, TNFa and IFNs.
 Pattern recognition receptors: functional characterization of TLRs
 Prostaglandins: role of prostaglandins, (PGE2, PGD2 and PGJ2) in the regulation of inflammation 
 Phagocytes: role of granulocytes and macrophages in the immune response 
 Gut immunity: functional characterization of mast cells
 Reproductive immunology: role of leukocytes and cytokines in spermatogenesis


 Development of oral immunostimulants and adjuvants
 Development of oral vaccines


 TNFR signaling in hematopoiesis, inflammation and cancer
 In vivo imaging of inflammation and tumour development
 Identification of new pharmacological compounds to treat chronic inflammation and cancer by high throughput screening 

Go to index



Dr. Isabel Cabas

 Role of GPER1 in fish immunity

Dr. Jorge Galindo-Villegas

 Oral immunostimulation and vaccination
 Interactions between microbiota and immunity 
Dr. Diana Garcia-Moreno 
 Epigenetic regulation of inflammation in zebrafish
 Implementation of site directed mutagenesis by Zinc-Finger nucleases

Dr. Han Mingming 

 Identification and functional characterization of new inflammasome components

Dr. Sylwia Tyrkalska

 Identification and functional characterization of new inflammasome components

PhD students

Elena Gomez

 Modelling the impact of inflammation on melanoma development using the zebrafish.

Nuria E. Gomez-Gonzalez

 Functional characterization of fish mast cells and role of histamine in innate and adaptive immunity.

Victoria Gomez-Abellan

 Role of prostaglandins in the regulation of fish immunity 

Fatma Hermi  

 Phylogenetic, molecular and functional characterization of fish prostaglandin receptors

Amanda E. Lopez-Canovas

 Slaughter stress in seabream muscle quality

Francisco J. Martinez-Morcillo

 Identification and validation  of new therapeutical targets and prognosis markers for psoriasis.

Francisco J. Martinez-Navarro

 Identification and validation  of new therapeutical targets and prognosis markers for psoriasis.

Maricarmen Rodenas

 Effects of endocrine disruptors on immunity

Ana Valera

 Identification and functional characterization of new inflammasome components


Inmaculada Fuentes

 Lab manager

Pedro Martinez

 Zebrafish facility manager


Former lab's members

Dr. Pablo Pelegrin

Unidad de Cirugia Experimental, Hospital Universitario Virgen de la Arrixaca. 30120 El Palmar. Murcia. Spain.

Dr. Jesus Garcia-Castillo

Grupo de Telómeros y Telomerasa. Unidad de Cirugía Experimental, IMIB-Arrixaca, carretera del Palmar, s/n. 30120 El Palmar, Murcia, Spain.

Dr. Gloria Lopez-Castejon

Faculty of Life Sciences, Michael Smith Building, Oxford Road, Manchester M13 9PT, UK

Dr. Elena Chaves-Pozo  

Centro Oceanográfico de Murcia (IEO),Mazarrón.Spain.

Dr. Ivan Mulero

Dr. Jaya Kumari

Dr. Francisco J. Roca

Department of Medicine, University of Cambridge, Cambridge, UK.

Dr. Patricia Castillo-Briceño

Institut de Biologie du Développement de Marseille Luminy, UMR 6216, CNRS-Université de la Méditerranée, Case 907, 13288 Marseille Cedex 09, France.

Dr. Iciar Muñoz

Dr. Sergio Liarte

Nicolas Gonzalez-Tejero

Ana Montalban

Zentrum für Medizinische Forschung (ZMF),Stiftingtalstraße 24, A-8010 Graz, Austria

Dr. Raquel Espin

Department of Cellular and Molecular Medicine and Section of Cell and Developmental Biology, University of California at San Diego, La Jolla, California, USA.

Dr. Diego Angosto

Dr. Erick Garcia-Garcia

Dr. Sofia de Oliveira 

Departments of Pediatrics and Medical Microbiology and Immunology, University of Wisconsin-Madison, Madison, WI 53706

Dr. Azucena Lopez-Muñoz

Dr. Jana Montero

Dr. Sergio Candel

Department of Medicine, University of Cambridge, Cambridge, UK.

Dr. Chiara Gabellini  


Go to index

Selected Publications (Click here for a full list)

MULERO I, SEPULCRE MP, MESEGUER J, GARCIA-AYALA A, MULERO V (2007). Histamine is stored in mast cells of most evolutionarily advanced fish and regulates the fish inflammatory response. Proc Natl Acad Sci USA. 104: 19434-19439. Full text

ROCA FJ, MULERO I, LOPEZ-MUÑOZ A, SEPULCRE MP, MESEGUER J, MULERO V (2008). Evolution of the inflammatory response in vertebrates: fish TNFa is a powerful activator of endothelial cells but hardly activates phagocytes. J Immunol 181:5071-5081. Full text

ALCARAZ-PEREZ F, MULERO V, CAYUELA ML (2008). Application of the Dual-Luciferase Reporter Assay to the Analysis of Promoter Activity in Zebrafish Embryos. BMC Biotechnol 8:81. Full text

SEPULCRE MP, ALCARAZ-PÉREZ F, LÓPEZ-MUÑOZ A, ROCA FJ, MESEGUER J, CAYUELA ML, MULERO V (2009). Evolution of LPS Recognition and Signaling: Fish TLR4 Does Not Recognize LPS and Negatively Regulates NF-kB Activation. J Immunol. 182: 1836-1845 Full text

LÓPEZ-MUÑOZ A, ROCA FJ, MESEGUER J, MULERO V (2009). New insights into the evolution of IFNs: Zebrafish group II IFNs induce a rapid and transient expression of IFN-dependent genes and display powerful antiviral activities. J Immunol 182: 3440-3449. Full text

OLAVARRÍA VH, SEPULCRE MP, FIGUEROA JE, MULERO V. (2010). Prolactin-induced production of ROS and IL-1b in leukocytes from the bony fish gilthead seabream involves Jak/Stat and NF-kB signaling pathways. J Immunol. 185:3873-3883. Full text

LÓPEZ-MUÑOZ A SEPULCRE MP, ROCA FJ, FIGUERAS A, MESEGUER J, MULERO V (2011). Evolutionary conserved pro-inflammatory and antigen presentation functions of zebrafish IFNγ revealed by transcriptomic and functional analysis. Mol. Immunol. 48:1073-1083. Full text

MULERO V, SEPULCRE MP, RAINGER GE, BUCKLEY CD (2011). Neutrophils live on a two way street. J Leukoc Biol 89:645-647. Full text.

OLAVARRÍA VH, FIGUEROA JE, MULERO V. (2012). Prolactin-induced activation of phagocyte NADPH oxidase in the teleost fish gilthead seabream involves the phosphorylation of p47phox by protein kinase C. Dev Comp Immunol. 36:216-221. Full text.

CABAS I, LIARTE S, GARCÍA-ALCÁZAR A, MESEGUER J, MULERO V, GARCÍA-AYALA A. (2012). 17α-Ethynylestradiol alters the immune response of the teleost gilthead seabream (Sparus aurata L.) both in vivo and in vitro. Dev Comp Immunol. 36:547-556 Full text.

LÓPEZ-MUÑOZ A SEPULCRE MP, GARCÍA-MORENO D, SEPULCRE MP, FUENTES I, BÉJAR J, MANCHADO M, ALVAREZ MC, MESEGUER J, MULERO V (2012). Viral nervous necrosis virus persistently replicates in the central nervous system of asymptomatic gilthead seabream and promotes a transient inflammatory response followed by the infiltration of IgM+ B lymphocytes. Dev Comp Immunol 7:429-437. Full text.

ANGOSTO D, LÓPEZ-CASTEJÓN G, LÓPEZ-MUÑOZ A SEPULCRE MP, ARIZCUN M, MESEGUER J, MULERO V (2012). Evolution of inflammasome functions in vertebrates: Inflammasome and caspase-1 trigger fish macrophage cell death but are dispensable for the processing of IL-1β. Innate Immun 18:815-824. Full text.

AGUILA S, CASTILLO-BRICEÑO P, SÁNCHEZ M, CABAS I, GARCÍA-ALCÁZAR A, MESEGUER J, MULERO V, GARCÍA-AYALA A. (2012). Specific and non-overlapping functions of testosterone and 11-ketotestosterone in the regulation of professional phagocyte responses in the teleost fish gilthead seabream. Mol Immunol. 53:218-226. Full text.

GALINDO-VILLEGAS J, GARCÍA-MORENO D, DE OLIVERIA S, MESEGUER J, MULERO V (2012). Regulation of immunity and disease resistance by commensal microbes and chromatin modifications during zebrafish development. Proc Natl Acad Sci USA 109: E2605-E2614. Full text. Full text Author Summary.

COMPAN V, BAROJA-MAZO A, LÓPEZ-CASTEJÓN G, GOMEZ AI, MARTÍNEZ CM, ANGOSTO D, MONTERO MT, HERRANZ AS, BAZÁN E, REIMERS D, MULERO V, PELEGRÍN P (2012). Cell volume regulation modulates NLRP3 inflammasome activation. Immunity 37:487-500. Full text.

ESPÍN R, ROCA FJ, CANDEL S, SEPULCRE MP, GONZÁLEZ-ROSA JM, ALCARAZ-PÉREZ F, MESEGUER J, CAYUELA ML, MERCADER N, MULERO V (2013). TNF receptors regulate vascular homeostasis through a caspase-8, caspase-2 and P53 apoptotic program that bypasses caspase-3. Dis Model Mech 6:383-396. Full text.

ANGOSTO D, LÓPEZ-CASTEJÓN G, MULERO V, PELEGRÍN P (2013). Response to Boyle et al.: Cell Swelling and the NLRP3 Inflammasome. Immunity 38:400-401. Full text.

DE OLIVEIRA S, REYES-ALDASORO CC, CANDEL S, RENSHAW SA, MULERO V, CALADO A. (2013). Cxcl8 (IL-8) mediates neutrophil recruitment and behavior in the zebrafish inflammatory response. J Immunol 190:4349-4359. Full text.

OLAVARRÍA VH, FIGUEROA JE, MULERO V (2013). Induction of genes encoding NADPH oxidase components and activation of interferon regulatory factor-1 by prolactin in fish macrophages. Innate Immun. 19:644-654. Full text.

GARCÍA-GARCIA E, GALINDO-VILLEGAS J, MULERO V. (2013). Mucosal immunity in the gut: the non-vertebrate perspective. Dev Comp Immunol 40:278-288. Full text.

MUÑOZ I, SEPULCRE MP, MESEGUER J, MULERO V (2013). Molecular cloning, phylogenetic analysis and functional characterization of soluble Toll-like receptor 5 in gilthead seabream, Sparus aurata. Fish Shellfish Immunol 35:36-45. Full text.

ANCHELIN M, ALCARAZ-PÉREZ F, MARTÍNEZ CM, BERNABÉ-GARCÍA M, MULERO V, CAYUELA ML (2013). Premature aging in telomerase-deficient zebrafish. Dis Model Mech. 6:1101-1112. Full text.

GALINDO-VILLEGAS J, MULERO I, GARCÍA-ALCAZAR A, MUÑOZ I, PEÑALVER-MELLADO M, STREITENBERGER S, SCAPIGLIATI G, MESEGUER J, MULERO V. (2013) Recombinant TNFα as oral vaccine adjuvant protects European sea bass against vibriosis: Insights into the role of the CCL25/CCR9 axis. Fish Shellfish Immunol 35:1260-1271. Full text.

ANGOSTO D, MONTERO J, LÓPEZ-MUÑOZ A, ALCARAZ-PÉREZ F, BIRD S, SARROPOULOU E, ABELLÁN E, MESEGUER J, SEPULCRE MP, MULERO V (2013). Identification and functional characterization of a new IL-1 family member, IL-1Fm2, in most evolutionarily advanced fish.. Innate Immun (in press). Full text.

CABAS I, RODENAS MC, ABELLÁN E, MESEGUER J, MULERO V, GARCÍA-AYALA A. (2013). Estrogen signaling through the g protein-coupled estrogen receptor regulates granulocyte activation in fish. J Immunol 191:4628-4639. Full text.

CABAS I, CHAVES-POZO E, GARCÍA-ALCÁZAR A, MESEGUER J, MULERO V, GARCÍA-AYALA A (2013). The effect of 17α-ethynylestradiol on steroidogenesis and gonadal cytokine gene expression is related to the reproductive stage in marine hermaphrodite fish. Mar Drugs. 11:4973-4992. Full text.

MUÑOZ I, SEPULCRE MP, MESEGUER J, MULERO V. (2014). Toll-like receptor 22 of gilthead seabream, Sparus aurata: Molecular cloning, expression profiles and post-transcriptional regulation. Dev Comp Immunol. 44:173-179. Full text.

SÁNCHEZ-HERNÁNDEZ M, ARIZCUN M, GARCÍA-ALCÁZAR A, SARROPOULOU E, MULERO V, GARCÍA-AYALA A. (2014). Fish granulocytes express a constitutively active androgen receptor variant. Dev Comp Immunol. 45:115-122. Full text.

ALCARAZ-PEREZ F, GARCIA-CASTILLO J, GARCIA-MORENO D, LOPEZ-MUNOZ A, ANCHELIN M, ANGOSTO D, ZON LI, MULERO V*, CAYUELA ML* (2014). A non-canonical function of telomerase RNA in the regulation of developmental myelopoiesis in zebrafish". Nat Commun 5:3228. Full text.

CANDEL S, DE OLIVEIRA S, LÓPEZ-MUÑOZ A, GARCÍA-MORENO D, ESPÍN-PALAZÓN R, TYRKALSKA SD, CAYUELA ML, RENSHAW SA, CORBALÁN R, VIDAL-ABARCA I, TSAI H-J, MESEGUER J, SEPULCRE MP, MULERO V. (2014). Tnfa Signaling Through Tnfr2 Protects Skin Against Oxidative Stress-Induced Inflammation. Plos Biol 12: e1001855. Full text.

DE OLIVEIRA S, LÓPEZ-MUÑOZ, A, CANDEL S, PELEGRIN P, CALADO A. MULERO V (2014). ATP modulates acute inflammation in vivo through dual oxidase 1-derived H2O2 production and NF-κB activation. J Immunol 192:5710-5719. Full text

GARCÍA-GARCÍA E, GÓMEZ-GONZÁLEZ NE, MESEGUER J, GARCÍA-AYALA A, MULERO V. (2014). Histamine regulates the inflammatory response of the tunicate Styela plicata. Dev Comp Immunol. 2014 46:382-391. Full text

MONTERO J, GÓMEZ-CASADO E, GARCÍA-ALCÁZAR A, MESEGUER J, MULERO V. (2014). Flagellin from Marinobacter algicola and Vibrio vulnificus activates the innate immune response of gilthead seabream. Dev Comp Immunol. 2014 47:160-167. Full text

GÓMEZ-GONZÁLEZ NE, GARCÍA-GARCÍA E, MONTERO J, GARCÍA-ALCÁZAR A, MESEGUER J, GARCÍA-AYALA A, MULERO V. (2014). Isolation of mast cells from the peritoneal exudate of the teleost fish gilthead sea bream (Sparus aurata L.). Fish Shellfish Immunol. 40:225-232. Full text

ESPÍN-PALAZÓN R, STACHURA DL, CAMPBELL CA, GARCÍA-MORENO D, DEL CID N, KIM AD, CANDEL S, MESEGUER J, MULERO V, TRAVER D. (2014). Proinflammatory signaling regulates hematopoietic stem cell emergence. Cell 159:1070-1085. Full text

DE OLIVEIRA S, LÓPEZ-MUÑOZ, A, MARTÍNEZ-NAVARRO FJ, GALINDO-VILLEGAS J, MULERO V, CALADO A (2014). Cxcl8-l1 and Cxcl8-l2 are required in the zebrafish defense against Salmonella Typhimurium. Dev Comp Immunol 49:44-48. Full text

GONZÁLEZ-STEGMAIER R, ROMERO A, ESTEPA A, MONTERO J, MULERO V, MERCADO L. (2015). Effects of recombinant flagellin B and its ND1 domain from Vibrio anguillarum on macrophages from gilthead seabream (Sparus aurata L.) and rainbow trout (Oncorhynchus mykiss, W.). Fish Shellfish Immunol 42:144-152. Full text

DE OLIVEIRA S, BOUDINOT P, CALADO A, MULERO V (2015). Duox1-Derived H2O2 Modulates Cxcl8 Expression and Neutrophil Recruitment via JNK/c-JUN/AP-1 Signaling and Chromatin Modifications. J Immunol 194:1523-1533. Full text

CANDEL S, SEPULCRE MP, ESPÍN-PALAZÓN R, TYRKALSKA SD, DE OLIVEIRA S, MESEGUER J, MULERO V (2015). Md1 and Rp105 regulate innate immunity and viral resistance in zebrafish. Dev Comp Immunol. 50:155-165. Full text

GÓMEZ-ABELLÁN V, MONTERO J, LÓPEZ-MUÑOZ A, FIGUERAS A, ARIZCUN M, MULERO V, SEPULCRE MP. (2015). Professional phagocytic granulocyte-derived PGD2 regulates the resolution of inflammation in fish. Dev Comp Immunol 52:182-191. Full text

LÓPEZ-MUÑOZ A, LIARTE S, GÓMEZ-GONZÁLEZ NE, CABAS I, MESEGUER J, GARCÍA-AYALA A, MULERO V. (2015). Estrogen receptor 2b deficiency impairs the antiviral response of zebrafish. Dev Comp Immunol 53:55-62. Full text

MONTALBAN-ARQUES A, DE SCHRYVER P, BOSSIER P, GORKIEWICZ G, MULERO V, GATLIN DM 3RD, GALINDO-VILLEGAS J. (2015) Selective Manipulation of the Gut Microbiota Improves Immune Status in Vertebrates. Front Immunol. 6:512. Full text

GONZÁLEZ-STEGMAIER R, GUZMÁN F, ALBERICIO F, VILLARROEL-ESPÍNDOLA F, ROMERO A, MULERO V, MERCADO L. (2015) A synthetic peptide derived from the D1 domain of flagellin induced the expression of proinflammatory cytokines in fish macrophages. Fish Shellfish Immunol. 47:239-244. Full text

RODENAS MC, CABAS I, ABELLÁN E, MESEGUER J, MULERO V, GARCÍA-AYALA A. (2015). Tamoxifen persistently disrupts the humoral adaptive immune response of gilthead seabream (Sparus aurata L.). Dev Comp Immunol. 53:283-292. Full text

GALINDO-VILLEGAS J, MONTALBAN-ARQUES A, LIARTE S, DE OLIVEIRA S, PARDO-PASTOR C, RUBIO-MOSCARDO F, MESEGUER J, VALVERDE MA, MULERO V (2016). Trpv4-mediated detection of hyposmotic stress by skin keratinocytes activates developmental immunity. J Immunol 196:738-749. Full text

TYRKALSKA SD, CANDEL S, ANGOSTO D, GÓMEZ-ABELLÁN V, MARTÍN-SÁNCHEZ F, GARCÍA-MORENO D, ZAPATA-PÉREZ R, SÁNCHEZ-FERRER A, SEPULCRE MP, PELEGRÍN P, MULERO V. (2016). Neutrophils mediate Salmonella Typhimurium clearance through the GBP4 inflammasome-dependent production of prostaglandins. Nat Commun (accepted 6 May 2016). Full text

ESPIN-PALAZON E, MARTÍNEZ-LÓPEZ A, ROCA FJ, LÓPEZ-MUÑOZ A, TYRKALSKA SD, CANDEL S, GARCÍA-MORENO D, FALCO A, MESEGUER J, ESTEPA A, MULERO V. (2016). TNFα impairs rhabdoviral clearance by inhibiting the host autophagic antiviral response. Plos Pathogens (accepted 20 May 2016). Full text

MONTERO J, GÓMEZ-ABELLÁN V, ARIZCUN M, MULERO V, SEPULCRE MP (2016). Prostaglandin E2 promotes M2 polarization of macrophages via a cAMP/CREB signaling pathway and deactivates granulocytes in teleost fish. Fish Shellfish Immunol. pii: S1050-4648(16)30404-1. Full text

CANDEL S, TYRKALSKA SD, GARCIA-MORENO D, MESEGUER J, MULERO V (2016). Identification of evolutionarily conserved Md1 splice variants that regulate innate immunity through differential induction of NF-кB. J Immunol. (accepted 13 June 2016).

Research grants

Our research is funded by:


Ministerio de Economía y Competitividad - Gobierno de España

Secretaría de Estado de Investigación, Desarrollo e Innovación - Ministerio de Economía y Competitividad - Gobierno de España



Seneca logo


Go to index

Positions available

No positions are available at this moment

Please contact us:

Go to index

Updated: June 2016