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The last 5 years have seen a remarkable extension of the innate immune system starting with what seemed at the time primarily cells that exert their influence during ontogeny of the immune system. So-called lymphoid tissue inducer (Lti) cells were soon joined by RORgt expressing innate lymphoid cells (ILC) that shaped immune responses beyond the determination of lymphoid architecture (Sawa et al., 2010). The latest addition to the growing family of ILC are characterized by the production of T helper (Th) 2-type cytokines even in the absence of the adaptive immune system (Moro et al., 2010; Neill et al., 2010; Price et al., 2010; Saenz et al., 2010).  T helper 2-type cytokine responses play a major role in the immunity to parasitic helminth infections, but are also implicated in the development of allergic diseases such as asthma (Paul and Zhu, 2010). The hallmark of Th2 mediated immune responses is the activation of mast cells, eosinophils, basophils, and goblet cell hyperplasia mediated by the cytokines IL-5, IL-9, and IL-13. Until recently, T cells of the Th cell type 2 (Th2) subset were thought to be the most abundant and important source of these cytokines. However, it became clear that a group of innate cells exist, that lack conventional markers typical for main lymphoid cell types and are able to produce high amounts of Th2-type cytokines (Fort et al., 2001; Hurst et al., 2002; Fallon et al., 2006). Such responses were shown to be dependent on the epithelial cell derived cytokine IL-25. More recently those ILC populations were further characterized by four independent research groups and called natural helper cells (NHCs; Moro et al., 2010), nuocytes (Neill et al., 2010), innate helper type 2 (Ih2) cells (Price et al., 2010), or multipotent progenitors (MPPs; Saenz et al., 2010). Nuocytes and MPPs reside in mesenteric lymph nodes and spleen, NHCs are found in the fat-associated lymphoid tissue and Ih2 cells are dispersed throughout the body, with most Ih2 cells recovered from the liver. All studies in support of earlier work established an important role for ILC as IL-25 but also IL-33 dependent cells in the context of helminth infections. In a recent review those newly discovered cells were classified as innate helper cells type 2 (ILC2; Spits and Di Santo, 2011) compared with RORγt expressing ILC subsets linked to cytokines such as IL-17 and IL-22 (ILC17 and ILC22). Lately, our own studies using an IL-9 fate reporter mouse model established that besides the Th2-type cytokines IL-5, IL-6, and IL-13 ILC2 also produce IL-9 (Wilhelm et al., 2011). However unlike IL-5 and IL-13 production in ILC, IL-9 is not permanently expressed but characterized by a very transient IL-2 dependent expression profile thus extending the portfolio of cytokines released by ILC.

Innate Lymphoid Cells and Type 2 (Th2) Mediated Immune Responses ? Pathogenic or Beneficial?