title: The laminated layer: Recent advances and insights into Echinococcus biology and evolution.
abstract: The laminated layer is the unique mucin-based extracellular matrix that protects Echinococcus larvae, and thus to an important extent, shapes host-parasite relationships in the larval echinococcoses. In 2011, we published twin reviews summarizing what was known about this structure. Since then, important advances have been made. Complete genomes and some RNAseq data are now available for E. multilocularis and E. granulosus, leading to the inference that the E. multilocularis LL is probably formed by a single type of mucin backbone, while a second apomucin subfamily additionally contributes to the E. granulosus LL. Previously suspected differences between E. granulosus and E. multilocularis in mucin glycan size have been confirmed and pinned down to the virtual absence of Galβ1-3 chains in E. multilocularis. The LL carbohydrates from both species have been found to interact selectively with the Kupffer cell receptor expressed in rodent liver macrophages, highlighting the ancestral adaptations to rodents as intermediate hosts and to the liver as infection site. Finally, LL particles have been shown to possess carbohydrate-independent mechanisms profoundly conditioning non-liver-specific dendritic cells and macrophages. These advances are discussed in an integrated way, and in the context of the newly determined phylogeny of Echinococcus and its taenid relatives.
authors: Cátedra de Inmunología, Departamento de Biociencias (Facultad de Química) e Instituto de Química Biológica (Facultad de Ciencias), Universidad de la República, Montevideo, Uruguay. Electronic address: adiaz@fq.edu.uy.
authors: Álvaro
authors: Á
authors: Díaz
authors: Cátedra de Inmunología, Departamento de Biociencias (Facultad de Química) e Instituto de Química Biológica (Facultad de Ciencias), Universidad de la República, Montevideo, Uruguay.
authors: Cecilia
authors: C
authors: Fernández
authors: Cátedra de Inmunología, Departamento de Biociencias (Facultad de Química) e Instituto de Química Biológica (Facultad de Ciencias), Universidad de la República, Montevideo, Uruguay.
authors: Álvaro
authors: Á
authors: Pittini
authors: Cátedra de Inmunología, Departamento de Biociencias (Facultad de Química) e Instituto de Química Biológica (Facultad de Ciencias), Universidad de la República, Montevideo, Uruguay.
authors: Paula I
authors: PI
authors: Seoane
authors: Institute of Immunology and Infection Research, Centre for Immunity, Infection and Evolution, School of Biological Sciences, University of Edinburgh, Edinburgh, UK.
authors: Judith E
authors: JE
authors: Allen
authors: Cátedra de Inmunología, Departamento de Biociencias (Facultad de Química) e Instituto de Química Biológica (Facultad de Ciencias), Universidad de la República, Montevideo, Uruguay.
authors: Cecilia
authors: C
authors: Casaravilla
mesh_terms: D000818:Animals; D005075:Biological Evolution; D002240:Carbohydrate Sequence; D004446:Echinococcus / Q000737:chemistry / Q000235:genetics / Q000276:immunology / Q000502:physiology*; D005752:Gastric Mucins / Q000737:chemistry / Q000235:genetics; D054794:Glycomics; D007113:Immunity, Innate; D009077:Mucins / Q000737:chemistry* / Q000235:genetics / Q000276:immunology; D011134:Polysaccharides / Q000737:chemistry / Q000235:genetics
chemical_list: D005752:Gastric Mucins; D009077:Mucins; D011134:Polysaccharides; C053366:apomucin
keywords: Carbohydrate; Echinococcus; Laminated layer; Lectin; Liver; Mucin