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in Rainbow Trout Oncorhynchus mykiss: The First Th1-Type Cytokine Characterized Functionally in Fish1
* Scottish Fish Immunology Research Centre, University of Aberdeen, Aberdeen, Scotland, United Kingdom;
Inland Station/National Research Institute of Aquaculture, Fisheries Research Agency, Tamaki, Mie, Japan;
Department of Biology, University of Waterloo, Waterloo, Ontario, Canada; and
Fisheries Research Services, Marine Laboratory, Aberdeen, United Kingdom
IFN-
is one of the key cytokines in defining Th1 immune responses. In this study, an IFN- homologue has been identified in rainbow trout Oncorhynchus mykiss, and its biological activities have been characterized. The trout IFN- cDNA is 1034 bp in length and translates into a 180-aa protein. The first intron of the trout IFN- gene contains highly polymorphic GACA minisatellites and 44-bp DNA repeats, giving rise to at least six alleles. IFN- is structurally conserved among vertebrates, and a signature motif has been identified. A nuclear localization sequence known to be crucial for IFN- biological activities is also present in the C-terminal region of the trout IFN-. The IFN- expression was induced in head kidney leukocytes by stimulation with PHA or poly(I:C) and in kidney and spleen of fish injected with poly(I:C). rIFN- produced in Escherichia coli significantly stimulated gene expression of IFN--inducible protein 10 (IP-10), MHC class II 
-chain, and STAT1, and enhanced respiratory burst activity in macrophages. Deletion of 29-aa residues from the C terminus containing the nuclear localization sequence motif resulted in loss of activity with respect to induction of IP-10 in RTS-11 cells. Moreover, IFN--induced IP-10 expression was completely abolished by the protein kinase C inhibitor staurosporine, and partially reduced by U0126, a specific inhibitor for ERKs. Taken together, the present study has demonstrated for the first time a functional IFN- homologue in a fish species, strongly suggesting a conserved Th1 immune response is most likely present in lower vertebrates.
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