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Untied States Department of Agriculture-Agriculture Research Service, National Center for Cool and Cold Water Aquaculture, Kearneysville, WV 25430
The TNF superfamily (TNFSF) of proteins are cytokines involved in diverse immunological and developmental pathways. Little is known about their evolution or expression in lower vertebrate species. Bioinformatic searches of Zebrafish, Tetraodon, and Fugu genome and other teleost expressed sequence tag databases identified 44 novel gene sequences containing a TNF homology domain. This work reveals the following: 1) teleosts possess orthologs of BAFF, APRIL, EDA, TWEAK, 4-1BBL, Fas ligand, LIGHT, CD40L, RANKL, and possibly TL1A; 2) the BAFF-APRIL subfamily is enriched by a third member, BALM, unique to fish; 3) orthologs of lymphotoxins 
and 
were not clearly identified in teleosts and are substituted by a related ligand, TNF-New; 4) as many as four TRAIL-like genes are present in teleosts, as compared with only one in mammals; and 5) T cell activation ligands OX40L, CD27L, CD30L, and GITRL were not identified in any fish species. Finally, we characterize mRNA expression of TNFSF members CD40L, LIGHT, BALM, APRIL, Fas ligand, RANKL, TRAIL-like, and TNF-New in rainbow trout, Oncorhynchus mykiss, immune and nonimmune tissues. In conclusion, we identified a total of 14 distinct TNFSF members in fishes, indicating expansion of this superfamily before the divergence of bony fish and tetrapods, 
360–450 million years ago. Based on these findings, we extend a model of TNFSF evolution and the coemergence of the vertebrate adaptive immune system.
The costs of publication of this article were defrayed in part by the payment of page charges. This article must therefore be hereby marked advertisement in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
1 This research was supported by Agricultural Research Service CRIS Project 1930-32000-002 "Host-Pathogen and Environmental Interactions in Cool and Cold Water Aquaculture."
2 Current address: United States Fish and Wildlife Service, Northeast Fishery Center-Fish Health Center, P.O. Box 155, Lamar, PA 16848.
3 Address correspondence and reprint requests to Dr. Gregory D. Wiens, National Center for Cool and Cold Water Aquaculture, 11861 Leetown Road, Kearneysville, WV 25430. E-mail address: greg.wiens{at}ars.usda.gov
4 Abbreviations used in this paper: TNFSF, tumor necrosis factor superfamily; AK, anterior kidney; APRIL, a proliferation-inducing ligand; BAFF, B cell-activating factor; BALM, BAFF-APRIL-like molecule; EDA, ectodysplasin; EST, expressed sequence tag; FasL, Fas ligand; GITRL, glucocorticoid-induced TNFR-related gene ligand; HOX, homeobox; LIGHT, lymphotoxin-like inducible protein that competes with glycoprotein D for binding herpesvirus entry mediator on T cell; LT, lymphotoxin; ORF, open reading frame; PK, posterior kidney; RANKL, receptor activator of NF-
B; THD, TNF homology domain; TL1A, TNF ligand-related molecule 1; TM, transmembrane; TMD, transmembrane domain; TRAIL, TNF-related apoptosis inducing ligand; TWEAK, TNF-like weak inducer of apoptosis; UTR, untranslated region.
5 The online version of this article contains supplemental material.
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