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      Colour variation in cichlid fish: Developmental mechanisms, selective pressures and evolutionary consequences

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          Abstract

          Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to complex social behaviour and morphological versatility, they are characterised by extensive diversity in colouration, both within and between species. Here, we review the cellular and molecular mechanisms underlying colour variation in this group and the selective pressures responsible for the observed variation. We specifically address the evidence for the hypothesis that divergence in colouration is associated with the evolution of reproductive isolation between lineages. While we conclude that cichlid colours are excellent models for understanding the role of animal communication in species divergence, we also identify taxonomic and methodological biases in the current research effort. We suggest that the integration of genomic approaches with ecological and behavioural studies, across the entire cichlid family and beyond it, will contribute to the utility of the cichlid model system for understanding the evolution of biological diversity.

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          Most cited references261

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          Signals, Signal Conditions, and the Direction of Evolution

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            Pleiotropy in the melanocortin system, coloration and behavioural syndromes.

            In vertebrates, melanin-based coloration is often associated with variation in physiological and behavioural traits. We propose that this association stems from pleiotropic effects of the genes regulating the synthesis of brown to black eumelanin. The most important regulators are the melanocortin 1 receptor and its ligands, the melanocortin agonists and the agouti-signalling protein antagonist. On the basis of the physiological and behavioural functions of the melanocortins, we predict five categories of traits correlated with melanin-based coloration. A review of the literature indeed reveals that, as predicted, darker wild vertebrates are more aggressive, sexually active and resistant to stress than lighter individuals. Pleiotropic effects of the melanocortins might thus account for the widespread covariance between melanin-based coloration and other phenotypic traits in vertebrates.
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              Genetics, development and evolution of adaptive pigmentation in vertebrates.

              The study of pigmentation has played an important role in the intersection of evolution, genetics, and developmental biology. Pigmentation's utility as a visible phenotypic marker has resulted in over 100 years of intense study of coat color mutations in laboratory mice, thereby creating an impressive list of candidate genes and an understanding of the developmental mechanisms responsible for the phenotypic effects. Variation in color and pigment patterning has also served as the focus of many classic studies of naturally occurring phenotypic variation in a wide variety of vertebrates, providing some of the most compelling cases for parallel and convergent evolution. Thus, the pigmentation model system holds much promise for understanding the nature of adaptation by linking genetic changes to variation in fitness-related traits. Here, I first discuss the historical role of pigmentation in genetics, development and evolutionary biology. I then discuss recent empirically based studies in vertebrates, which rely on these historical foundations to make connections between genotype and phenotype for ecologically important pigmentation traits. These studies provide insight into the evolutionary process by uncovering the genetic basis of adaptive traits and addressing such long-standing questions in evolutionary biology as (1) are adaptive changes predominantly caused by mutations in regulatory regions or coding regions? (2) is adaptation driven by the fixation of dominant mutations? and (3) to what extent are parallel phenotypic changes caused by similar genetic changes? It is clear that coloration has much to teach us about the molecular basis of organismal diversity, adaptation and the evolutionary process.
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                Author and article information

                Journal
                Semin Cell Dev Biol
                Semin. Cell Dev. Biol
                Seminars in Cell & Developmental Biology
                Academic Press
                1084-9521
                1096-3634
                1 June 2013
                June 2013
                : 24
                : 6-7
                : 516-528
                Affiliations
                [a ]University of Groningen, Behavioural Biology, PO Box 11103, 9700 CC Groningen, The Netherlands
                [b ]Institute of Zoology, University of Graz, Universitätsplatz 2, A-8010 Graz, Austria
                Author notes
                [* ]Corresponding author. Tel.: +31 0 50 3632196. m.e.maan@ 123456rug.nl
                [1]

                Tel.: +43 0 316 380 5601.

                Article
                S1084-9521(13)00069-4
                10.1016/j.semcdb.2013.05.003
                3778878
                23665150
                010f6d59-2c81-4d31-a5ef-6ee409eaeafa
                © 2013 The Authors

                This document may be redistributed and reused, subject to certain conditions.

                History
                Categories
                Review

                Developmental biology
                cichlidae,natural selection,pigmentation,polymorphism,sexual selection,speciation

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