The eosinophil is a granulocyte prominent in allergic diseases and inflammatory responses against helminthic parasites. The eosinophil was named by Paul Ehrlich in 1879, and derives from the intense staining of its granules with the acidic dye eosin. It has been the subject of extensive investigation ever since. It is strongly associated with human diseases involving mucosal surfaces, such as allergic asthma, atopic dermatitis and gastrointestinal disorders. Eosinophils are likely involved in tissue homeostasis, modulation of adaptive immune responses, innate immunity to certain microbes and pathological changes in allergic disorders. Thus, the eosinophil is considered a multifunctional leukocyte that contributes to a wide variety of physiological and pathological processes, depending on its location and activation status. Further studies will be necessary to better understand the biology of this extraordinary leukocyte and to reveal the importance of the cell in human health and disease.

Gleich GJ, Adolphson CR: The eosinophil- ic leukocyte: structure and function. Adv Immunol 1986;39:177-253.
Mishra A, Hogan SP, Lee JJ, Foster PS, Rothenberg ME: Fundamental signals that regulate eosinophil homing to the gastrointestinal tract. J Clin Invest 1999;103:1719-1727.
Molofsky AB, Nussbaum JC, Liang HE, Van Dyken SJ, Cheng LE, Mohapatra A, Chawla A, Locksley RM: Innate lymphoid type 2 cells sustain visceral adipose tissue eosinophils and alternatively activated macrophages. J Exp Med 2013, in press.
Robertson SA, Mau VJ, Hudson SN, Tremellen KP: Cytokine-leukocyte networks and the establishment of pregnancy. Am J Reprod Immunol 1997;37:438-442.
Gouon-Evans V, Rothenberg ME, Pollard JW: Postnatal mammary gland development requires macrophages and eosinophils. Development 2000;127:2269-2282.
Furuta GT, Nieuwenhuis EE, Karhausen J, Gleich G, Blumberg RS, Lee JJ, Ackerman SJ: Eosinophils alter colonic epithelial barrier function: role for major basic protein. Am J Physiol Gastrointest Liver Physiol 2005;289:G890-G897.
Yousefi S, Gold JA, Andina N, Lee JJ, Kelly AM, Kozlowski E, Schmid I, Straumann A, Reichenbach J, Gleich GJ, Simon HU: Catapult-like release of mitochondrial DNA by eosinophils contributes to antibacterial defense. Nat Med 2008;14:949-953.
Wu D, Molofsky AB, Liang HE, Ricardo-Genzalez RR, Jouihan HA, Bando JI, Chawla A, Locksley RM: Eosinophils sustain adipose alternatively activated macrophages associated with glucose homeostasis. Science 2011;332:243-249.
Throsby M, Herbelin A, Pleau JM, Dardenne M: CD11c+ eosinophils in the murine thymus: developmental regulation and recruitment upon MHC class I-restricted thymocyte deletion. J Immunol 2000;165:1965-1975.
Chu VT, Fröhlich A, Steinhauser G, Scheel T, Roch T, Fillatreau S, Lee JJ, Löhning M, Berek C: Eosinophils are required for the maintenance of plasma cells in the bone marrow. Nat Immunol 2011;12:151-159.
Hogan SP, Rosenberg HF, Moqbel R, Phipps S, Foster PS, Lacy P, Kay AB, Rothenberg ME: Eosinophils: biological properties and role in health and disease. Clin Exp Allergy 2008;38:709-750.
Moqbel R, Lacy P: New concepts in effector functions of eosinophil cytokines. Clin Exp Allergy 2000;30:1667-1671.
Kita H, Ohnishi T, Okubo Y, Weiler D, Abrams JS, Gleich GJ: Granulocyte/macrophage colony-stimulating factor and interleukin 3 release from human peripheral blood eosinophils and neutrophils. J Exp Med 1991;174:745-748.
Levi-Schaffer F, Garbuzenko E, Rubin A, Reich R, Pickholz D, Gillery P, Emonard H, Nagler A, Maquart FA: Human eosinophils regulate human lung- and skin-derived fibroblast properties in vitro: a role for transforming growth factor beta (TGF-beta). Proc Natl Acad Sci USA 1999;96:9660-9665.
Burgel PR, Lazarus SC, Tam DC, Ueki IF, Atabai K, Birch M, Nadel JA: Human eosinophils induce mucin production in airway epithelial cells via epidermal growth factor receptor activation. J Immunol 2001;167:5948-5954.
Kobayashi H, Gleich GJ, Butterfield JH, Kita H: Human eosinophils produce neurotrophins and secrete nerve growth factor on immunologic stimuli. Blood 2002;99:2214-2220.
Nonaka M, Nonaka R, Woolley K, Adelroth E, Miura K, Okhawara Y, et al: Distinct immunohistochemical localization of IL-4 in human inflamed airway tissues. IL-4 is localized to eosinophils in vivo and is released by peripheral blood eosinophils. J Immunol 1995;155:3234-3244.
Moqbel R, Ying S, Barkans J, Newman TM, Kimmitt P, Wakelin M, Taborda-Barata L, Meng Q, Corrigan CJ, Durham SR, Kay AB: Identification of messenger RNA for IL-4 in human eosinophils with granule localization and release of the translated product. J Immunol 1995;155:4939-4947.
Bandeira-Melo C, Sugiyama K, Woods LJ, Weller PF: Cutting edge: eotaxin elicits rapid vesicular transport-mediated release of preformed IL-4 from human eosinophils. J Immunol 2001;166:4813-4817.
Terness P, Bauer TM, Röse L, Dufter C, Watzlik A, Simon H, Opelz G: Inhibition of allogeneic T cell proliferation by indoleamine 2,3-dioxygenase-expressing dendritic cells: mediation of suppression by tryptophan metabolites. J Exp Med 2002;196:447-457.
Odemuyiwa SO, Ghahary A, Li Y, Puttagunta L, Lee JE, Musat-Marcu S, Ghahary A, Moqbel R: Cutting edge: human eosinophils regulate T cell subset selection through indoleamine 2,3-dioxygenase. J Immunol 2004;173:5909-5913.
Yang D, Rosenberg HF, Chen Q, Dyer KD, Kurosaka K, Oppenheim JJ: Eosinophil-derived neurotoxin (EDN), an antimicrobial protein with chemotactic activities for dendritic cells. Blood 2003;102:3396-3403.
Yang D, Chen Q, Rosenberg HF, Rybak SM, Newton DL, Wang ZY, Fu Q, Tchernev VT, Wang M, Schweitzer B, Kingsmore SF, Patel DD, Oppenheim JJ, Howard OM: Human ribonuclease A superfamily members, eosinophil-derived neurotoxin and pancreatic ribonuclease, induce dendritic cell maturation and activation. J Immunol 2004;173:6134-6142.
Yang D, Chen Q, Su SB, Zhang P, Kurosaka K, Caspi RR, Michalek SM, Rosenberg HF, Zhang N, Oppenheim JJ: Eosinophil-derived neurotoxin acts as an alarmin to activate the TLR2-MyD88 signal pathway in dendritic cells and enhances Th2 immune responses. J Exp Med 2008;205:79-90.
Spencer LA, Weller PF: Eosinophils and Th2 immunity: contemporary insights. Immunol Cell Biol 2010;88:250-256.
Voehringer D, Shinkai K, Locksley RM: Type 2 immunity reflects orchestrated recruitment of cells committed to IL-4 production. Immunity 2004;20:267-277.
Bartemes KR, Iijima K, Kobayashi T, Kephart GM, McKenzie AN, Kita H: IL-33-responsive lineage -CD25+CD44hi lymphoid cells mediate innate type-2 immunity and allergic inflammation in the lungs. J Immunol 2012;188:1503-1513.
Shinkai K, Mohrs M, Locksley RM: Helper T cells regulate type-2 innate immunity in vivo. Nature 2002;420:825-829.
Sabin EA, Pearce EJ: Early IL-4 production by non-CD4+ cells at the site of antigen deposition predicts the development of a T helper 2 cell response to Schistosoma mansoni eggs. J Immunol 1995;155:4844-4853.
Walker JA, Barlow JL, McKenzie AN: Innate lymphoid cells - how did we miss them? Nat Rev Immunol 2013;13:75-87.
Mattes J, Yang M, Mahalingam S, Kuehr J, Webb DC, Simson L, Hogan SP, Koskinen A, McKenzie AN, Dent LA, Rothenberg ME, Matthaei KI, Young IG, Foster PS: Intrinsic defect in T cell production of interleukin (IL)-13 in the absence of both IL-5 and eotaxin precludes the development of eosinophilia and airways hyperreactivity in experimental asthma. J Exp Med 2002;195:1433-1444.
Lee JJ, Dimina D, Macias MP, Ochkur SI, McGarry MP, O'Neill KR, Protheroe C, Pero R, Nguyen T, Cormier SA, Lenkiewicz E, Colbert D, Rinaldi L, Ackerman SJ, Irvin CG, Lee NA: Defining a link with asthma in mice congenitally deficient in eosinophils. Science 2004;305:1773-1776.
Yu C, Cantor AB, Yang H, Browne C, Wells RA, Fujiwara Y, Orkin SH: Targeted deletion of a high-affinity GATA-binding site in the GATA-1 promoter leads to selective loss of the eosinophil lineage in vivo. J Exp Med 2002;195:1387-1395.
Walsh ER, Sahu N, Kearley J, Benjamin E, Kang BH, Humbles A, August A: Strain-specific requirement for eosinophils in the recruitment of T cells to the lung during the development of allergic asthma. J Exp Med 2008;205:1285-1292.
Jacobsen EA, Ochkur SI, Pero RS, Taranova AG, Protheroe CA, Colbert DC, Lee NA, Lee JJ: Allergic pulmonary inflammation in mice is dependent on eosinophil-induced recruitment of effector T cells. J Exp Med 2008;205:699-710.
Flood-Page P, Menzies-Gow A, Phipps S, Ying S, Wangoo A, Ludwig MS, Barnes N, Robinson D, Kay AB: Anti-IL-5 treatment reduces deposition of ECM proteins in the bronchial subepithelial basement membrane of mild atopic asthmatics. J Clin Invest 2003;112:1029-1036.
Abu-Ghazaleh RI, Gleich GJ, Prendergast FG: Interaction of eosinophil granule major basic protein with synthetic lipid bilayers: a mechanism for toxicity. J Membr Biol 1992;128:153-164.
Rosenberg HF, Domachowske JB: Eosinophils, eosinophil ribonucleases, and their role in host defense against respiratory virus pathogens. J Leukoc Biol 2001;70:691-698.
Adamko DJ, Yost BL, Gleich GJ, Fryer AD, Jacoby DB: Ovalbumin sensitization changes the inflammatory response to subsequent parainfluenza infection. Eosinophils mediate airway hyperresponsiveness, m(2) muscarinic receptor dysfunction, and antiviral effects. J Exp Med 1999;190:1465-1478.
Kita H, Weiler DA, Abu-Ghazaleh R, Sanderson CJ, Gleich GJ: Release of granule proteins from eosinophils cultured with IL-5. J Immunol 1992;149:629-635.
Bracke M, Dubois GR, Bolt K, Bruijnzeel PL, Vaerman JP, Lammers JW, Koenderman L: Differential effects of the T helper cell type 2-derived cytokines IL-4 and IL-5 on ligand binding to IgG and IgA receptors expressed by human eosinophils. J Immunol 1997;159:1459-1465.
Valerius T, Repp R, Kalden JR, Platzer E: Effects of IFN on human eosinophils in comparison with other cytokines. A novel class of eosinophil activators with delayed onset of action. J Immunol 1990;145:2950-2958.
Saenz SA, Taylor BC, Artis D: Welcome to the neighborhood: epithelial cell-derived cytokines license innate and adaptive immune responses at mucosal sites. Immunol Rev 2008;226:172-190.
Wong CK, Hu S, Cheung PF, Lam CW: Thymic stromal lymphopoietin induces chemotactic and prosurvival effects in eosinophils: implications in allergic inflammation. Am J Respir Cell Mol Biol 2010;43:305-315.
Cherry WB, Yoon J, Bartemes KR, Iijima K, Kita H: A novel IL-1 family cytokine, IL-33, potently activates human eosinophils. J Allergy Clin Immunol 2008;121:1484-1490.
Nagase H, Okugawa S, Ota Y, Yamaguchi M, Tomizawa H, Matsushima K, Ohta K, Yamamoto K, Hirai K: Expression and function of Toll-like receptors in eosinophils: activation by Toll-like receptor 7 ligand. J Immunol 2003;171:3977-3982.
Miike S, McWilliam AS, Kita H: Trypsin induces activation and inflammatory mediator release from human eosinophils through protease-activated receptor-2. J Immunol 2001;167:6615-6622.
Miike S, Kita H: Human eosinophils are activated by cysteine proteases and release inflammatory mediators. J Allergy Clin Immunol 2003;111:704-713.
Matsuwaki Y, Wada K, White TA, Benson LM, Charlesworth MC, Checkel JL, Inoue Y, Hotta K, Ponikau JU, Lawrence CB, Kita H: Recognition of fungal protease activities induces cellular activation and eosinophil-derived neurotoxin release in human eosinophils. J Immunol 2009;183:6708-6716.
Wada K, Matsuwaki Y, Yoon J, Benson LM, Checkel JL, Bingemann TA, Kita H: Inflammatory responses of human eosinophils to cockroach are mediated through protease-dependent pathways. J Allergy Clin Immunol 2010;126:169-172.
Bush RK, Prochnau JJ: Alternaria-induced asthma. J Allergy Clin Immunol 2004;113:227-234.
Inoue Y, Matsuwaki Y, Shin SH, Ponikau JU, Kita H: Nonpathogenic, environmental fungi induce activation and degranulation of human eosinophils. J Immunol 2005;175:5439-5447.
Yoon J, Ponikau JU, Lawrence CB, Kita H: Innate antifungal immunity of human eo- sinophils mediated by a beta 2 integrin, CD11b. J Immunol 2008;181:2907-2915.
Bianchi ME: DAMPs, PAMPs and alarmins: all we need to know about danger. J Leukoc Biol 2007;81:1-5.
Kobayashi T, Kouzaki H, Kita H: Human eosinophils recognize endogenous danger signal crystalline uric acid and produce proinflammatory cytokines mediated by autocrine ATP. J Immunol 2010;184:6350-6358.
Lotfi R, Herzog GI, DeMarco RA, Beer-Stolz D, Lee JJ, Rubartelli A, Schrezenmeier H, Lotze MT: Eosinophils oxidize damage-associated molecular pattern molecules derived from stressed cells. J Immunol 2009;183:5023-5031.
Abu-Ghazaleh RI, Fujisawa T, Mestecky J, Kyle RA, Gleich GJ: IgA-induced eosinophil degranulation. J Immunol 1989;142:2393-2400.
Motegi Y, Kita H: Interaction with secretory component stimulates effector functions of human eosinophils but not of neutrophils. J Immunol 1998;161:4340-4346.
Kaneko M, Horie S, Kato M, Gleich GJ, Kita H: A crucial role for beta 2 integrin in the activation of eosinophils stimulated by IgG. J Immunol 1995;155:2631-2641.
Kato M, Kimura H, Motegi Y, Tachibana A, Minakami H, Morikawa A, Kita H: Platelet-activating factor activates two distinct effector pathways in human eosinophils. J Immunol 2002;169:5252-5259.
Horie S, Kita H: CD11b/CD18 (Mac-1) is required for degranulation of human eosinophils induced by human recombinant granulocyte-macrophage colony-stimulating factor and platelet-activating factor. J Immunol 1994;152:5457-5467.
Kato M, Abraham RT, Okada S, Kita H: Ligation of the beta2 integrin triggers activation and degranulation of human eosinophils. Am J Respir Cell Mol Biol 1998;18:675-686.
Nagata M, Sedgwick JB, Kita H, Busse WW: Granulocyte macrophage colony-stimulating factor augments ICAM-1 and VCAM-1 activation of eosinophil function. Am J Respir Cell Mol Biol 1998;19:158-166.
Yoon J, Terada A, Kita H: CD66b regulates adhesion and activation of human eosinophils. J Immunol 2007;179:8454-8462.
Mestas J, Hughes CC: Of mice and not men: differences between mouse and human immunology. J Immunol 2004;172:2731-2738.
Rosenberg HF: The eosinophil ribonucleases. Cell Mol Life Sci 1998;54:795-803.
Dyer KD, Percopo CM, Xie Z, Yang Z, Kim JD, Davoine F, Lacy P, Druey KM, Moqbel R, Rosenberg HF: Mouse and human eosinophils degranulate in response to platelet-activating factor (PAF) and lysoPAF via a PAF-receptor-independent mechanism: evidence for a novel receptor. J Immunol 2010;184:6327-6334.
Borchers MT, Ansay T, DeSalle R, Daugherty BL, Shen H, Metzger M, Lee NA, Lee JJ: In vitro assessment of chemokine receptor-ligand interactions mediating mouse eosinophil migration. J Leukoc Biol 2002;71:1033-1041.
Medzhitov R: Origin and physiological roles of inflammation. Nature 2008;454:428-435.
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