Trans-species polymorphism at antimicrobial innate immunity cathelicidin genes of Atlantic cod and related species

Institute of Life and Environmental Sciences, University of Iceland, Reykjavík, Iceland
DOI
10.7287/peerj.preprints.924v1
Subject Areas
Aquaculture, Fisheries and Fish Science, Evolutionary Studies, Genetics, Marine Biology, Immunology
Keywords
Atlantic cod, Innate immunity, Cathelicidin, Balancing selection, Trans-species polymorphism, Gadids
Copyright
© 2015 Halldórsdóttir et al.
Licence
This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ PrePrints) and either DOI or URL of the article must be cited.
Cite this article
Halldórsdóttir K, Árnason E. 2015. Trans-species polymorphism at antimicrobial innate immunity cathelicidin genes of Atlantic cod and related species. PeerJ PrePrints 3:e924v1

Abstract

Natural selection, the most important force in evolution, comes in three forms. Negative purifying selection removes deleterious variation and maintains adaptations. Positive directional selection fixes beneficial variants, producing new adaptations. Balancing selection maintains variation in a population. Important mechanisms of balancing selection include heterozygote advantage, frequency-dependent advantage of rarity, and local and fluctuating episodic selection. A rare pathogen gains an advantage because host defenses are predominantly effective against prevalent types. Similarly, a rare immune variant gives its host an advantage because the prevalent pathogens cannot escape the host's apostatic defense. Due to the stochastic nature of evolution, neutral variation may accumulate on genealogical branches, but trans-species polymorphisms are rare under neutrality and are strong evidence for balancing selection. Balanced polymorphism maintains diversity at the major histocompatibility complex (MHC) in vertebrates. The Atlantic cod is missing genes for both MHC-II and CD4, vital parts of the adaptive immune system. Nevertheless, cod are healthy in their ecological niche, maintaining large populations that support major commercial fisheries. Innate immunity is of interest from an evolutionary perspective, particularly in taxa lacking adaptive immunity. Here, we analyze extensive amino acid and nucleotide polymorphisms of the cathelicidin gene family in Atlantic cod and closely related taxa. There are three major clusters, Cath1, Cath2, and Cath3, that we consider to be paralogous genes. There is extensive nucleotide and amino acid allelic variation between and within clusters. The major feature of the results is that the variation clusters by alleles and not by species in phylogenetic trees and discriminant analysis of principal components. The three groups show trans-species polymorphism that is older than speciation and that is evidence for balancing selection maintaining the variation. Using Bayesian and likelihood methods positive and negative selection is evident at sites in the conserved part of the genes and, to a larger extent, in the active part which also shows episodic diversifying selection, further strengthening the argument for balancing selection.

Author Comment

This is a submission to PeerJ for review.

Supplemental Information

Supplementary data for the manuscript: Trans-species Polymorphism at Antimicrobial Innate Immunity Cathelicidin Genes of Atlantic cod and Related Species

DOI: 10.7287/peerj.preprints.924v1/supp-1