The vertebrate immune system is comprised of numerous distinct and interdependent components Every component has its own inherent protective value, and the final combination of them is likely to be related to an animal’s immunological history and evolutionary development Vertebrate immune system consists of both systemic and mucosal immune compartments, but it is the mucosal immune system which protects the body from the first encounter of pathogens According to anatomical location, the mucosa-associated lymphoid tissue, in teleost fish is subdivided into gut-, skin-, and gill-associated lymphoid tissue and most available studies focus on gut The purpose of this paper is to summarise the current knowledge of the immunological defences present in skin mucosa as a very important part of the fish immune system, serving as an anatomical and physiological barrier against external hazards Interest in defence mechanism of fish arises from a need to develop health management tools to support a growing finfish aquaculture industry, while at the same time addressing questions concerning origins and evolution of immunity in vertebrates Increased knowledge of fish mucosal immune system will facilitate the development of novel vaccination strategies in fish

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An Overview of the Immunological Defenses in Fish Skin

This article reviews the history of gyrodactylid research focussing on the unique anatomy, behaviour, ecology and evolution of the viviparous forms while identifying gaps in our knowledge and directions for future research. We provide the first summary of research on the oviparous gyrodactylids from South American catfish, and highlight the plesiomorphic characters shared by gyrodactylids and other primitive monogeneans. Of these, the most important are the crawling, unciliated larva and the spike sensilla of the cephalic lobes. These characters allow gyrodactylids to transfer between hosts at any stage of the life cycle, without a specific transmission stage. We emphasise the importance of progenesis in shaping the evolution of the viviparous genera and discuss the relative extent of progenesis in the different genera. The validity of the familial classification is discussed and we conclude that the most significant division within the family is between the oviparous and the viviparous genera. The older divisions into Isancistrinae and Polyclithrinae should be allowed to lapse. We discuss approaches to the taxonomy of gyrodactylids, and we emphasise the importance of adequate morphological and molecular data in new descriptions. Host specificity patterns in gyrodactylids are discussed extensively and we note the importance of host shifts, revealed by molecular data, in the evolution of gyrodactylids. To date, the most closely related gyrodactylids have not been found on closely related hosts, demonstrating the importance of host shifts in their evolution. The most closely related species pair is that of G. salaris and G. thymalli, and we provide an account of the patterns of evolution taking place in different mitochondrial clades of this species complex. The host specificity of these clades is reviewed, demonstrating that, although each clade has its preferred host, there is a range of specificity to different salmonids, providing opportunities for complex patterns of survival and interbreeding in Scandinavia. At the same time, we identify trends in systematics and phylogeny relevant to the G. salaris epidemics on Atlantic salmon in Norway, which can be applied more generally to parasite epidemiology and evolution. Although much of gyrodactylid research in the last 30 years has been directed towards salmonid parasites, there is great potential in using other experimental systems, such as the gyrodactylids of poeciliids and sticklebacks. We also highlight the role of glacial lakes and modified river systems during the ice ages in gyrodactylid speciation, and suggest that salmon infecting clades of G. salaris first arose from G. thymalli in such lakes, but failed to spread fully across Scandinavia before further dispersal was ended by rising sea levels. This dispersal has been continued by human activity, leading to the appearance of G. salaris as a pathogen in Norway. We review the history and current status of the epidemic, and current strategies for elimination of the parasite from Norway. Finally, we consider opportunities for further spread of the parasite within and beyond Europe.

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The biology of gyrodactylid monogeneans: the "Russian-doll killers".

The main objective of this study was to assess the effects of 4 g·kg-1dietary MOS inclusion in soybean oil (SBO) and fish oil (FO) based diets on the gut health and skin mucosa mucus production of European sea bass juveniles after 8 weeks of feeding. Dietary MOS, regardless of the oil source, promoted growth. The intestinal somatic index was not affected, however SBO reduced the intestinal fold length, while dietary MOS increased it. The dietary oil source fed produced changes on the posterior intestine fatty acid profiles irrespective of MOS dietary supplementation. SBO down-regulated the gene expression of TCRβ, COX2, IL-1β, TNFα, IL-8, IL-6, IL-10, TGFβ and Ig and up-regulated MHCII. MOS up-regulated the expression of MHCI, CD4, COX2, TNFα and Ig when included in FO based diets. However, there was a minor up-regulating effect on these genes when MOS was supplemented in the SBO based diet. Both dietary oil sources and MOS affected mean mucous cell areas within the posterior gut, however the addition of MOS to a SBO diet increased the mucous cell size over the values shown in FO fed fish. Dietary SBO also trends to reduce mucous cell density in the anterior gut relative to FO, suggesting a lower overall mucosal secretion. There are no effects of dietary oil or MOS in the skin mucosal patterns. Complete replacement of FO by SBO, modified the gut fatty acid profile, altered posterior gut GALT-related gene expression and gut mucous cells patterns, induced shorter intestinal folds and tended to reduce European sea bass growth. However, when combined with MOS, the harmful effects of SBO appear to be partially balanced by moderating the down-regulation of certain GALT-related genes involved in the functioning of gut mucous barrier and increasing posterior gut mucous cell diffusion rates, thus helping to preserve immune homeostasis. This denotes the importance of a balanced dietary n3/n6 ratio for an appropriate GALT-immune response against MOS in European sea bass.

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Dietary Mannan Oligosaccharides: Counteracting the Side Effects of Soybean Meal Oil Inclusion on European Sea Bass (Dicentrarchus labrax) Gut Health and Skin Mucosa Mucus Production?

The effects on rainbow trout, Oncorhynchus mykiss (Walbaum), immune parameters by differently formulated fish feed types containing immunostimulants have been tested in a double-blind, duplicated and controlled study performed over 50 days. A total of 800 rainbow trout (10-12 g) were kept in eight duplicate fish tanks (each containing 100 fish) and fed at a daily feeding rate of 1.5% of the biomass. The feed types were (1) control feed (C) without additives, (2) feed containing beta-glucan, nucleotides, manno-oligosaccharides (MOS), vitamins C and E (GNMCE), (3) feed containing probiotic bacteria and plant extracts (PP) and (4) feed with nucleotides, manno-oligosaccharides, vitamins C and E (NMCE). Plasma lysozyme activity was increased in fish fed two feed types (GNMCE and NMCE) but slightly depressed in fish fed PP. A non-significant trend for a higher mucous cell density at days 30 and 50 was shown in all fish receiving feeds with additives compared to the control group. All fish became infected with Ichthyophthirius multifiliis when exposed, but fish fed GNMCE showed a significantly lower infection both at days 30 and 50. Expression of genes encoding C3 and MHCII was significantly up-regulated in fish fed GNMCE for 50 days, and the expression of genes coding Hepcidin was significantly down-regulated in fish fed NMCE for 50 days. Beta-glucan was the single component, when used in combination with other feed ingredients, which was found associated with increased parasite resistance, increased lysozyme and immune gene up-regulation.

Comparative effects of four feed types on white spot disease susceptibility and skin immune parameters in rainbow trout, Oncorhynchus mykiss (Walbaum).

Mucous cell size and distribution were investigated in the skin of five salmon using a novel stereology-based methodology: one (48 cm) fish to test 15 tissue treatment combinations on measures of cell area and density on the dorsolateral region and, using the most suitable treatment, we mapped mucous cell differences between body regions on four (52 cm) salmon, comprising a male and a female on each of two diets. The section site, decalcification, embedding medium and plane of sectioning all impacted significantly on mucous cell size, whereas mucous cell density is more robust. There were highly significant differences in both mucosal density and mean mucous cell size depending on body site: the dorsolateral skin of the four salmon had significantly denser (about 8% of skin area) and larger (mean about 160 μm(2)) mucous cells, whereas the lowest mean density (about 4%) and smallest mean area (115 μm(2)) were found on the head. We found that 100 random measurements may be sufficient to distinguish differences >7 μm(2) in mean mucous cell areas. The results further suggest that salmon exhibit a dynamic repeatable pattern of mucous cell development influenced by sex, diet and possibly strain and season.

Body site matters: an evaluation and application of a novel histological methodology on the quantification of mucous cells in the skin of Atlantic salmon, Salmo salar L.

Atlantic salmon, Salmo salar, is normally rather refractive to infection with the ectoparasitic monogenean Gyrodactylus derjavini but dexamethasone treatment of the host increases the susceptibility. The causative mechanisms were elucidated in this work. Groups of Atlantic salmon were treated by intra-peritoneal dexamethasone injections and subsequently infected with G. derjavini. It was shown that both the infection level and the mucous cell density of caudal and pelvic fins were affected by the treatment. Significantly higher mucous cell densities were found on infected and treated fish whereas non-infected and treated fish showed no significant elevation of cell density. This suggests that mucous cell discharge elicited by infection is inhibited by the drug. The association with elevated parasite counts in these fish can be explained either by decreased anti-parasitic mucus action or by parasite predilection for intact mucous cells.

S.H. Olafsdottir

Papers

Dexamethasone treatment affects skin mucous cell density in Gyrodactylus derjavini infected Salmo salar.