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      A genomic comparison of two termites with different social complexity

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          Abstract

          The termites evolved eusociality and complex societies before the ants, but have been studied much less. The recent publication of the first two termite genomes provides a unique comparative opportunity, particularly because the sequenced termites represent opposite ends of the social complexity spectrum. Zootermopsis nevadensis has simple colonies with totipotent workers that can develop into all castes (dispersing reproductives, nest-inheriting replacement reproductives, and soldiers). In contrast, the fungus-growing termite Macrotermes natalensis belongs to the higher termites and has very large and complex societies with morphologically distinct castes that are life-time sterile. Here we compare key characteristics of genomic architecture, focusing on genes involved in communication, immune defenses, mating biology and symbiosis that were likely important in termite social evolution. We discuss these in relation to what is known about these genes in the ants and outline hypothesis for further testing.

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          Most cited references61

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          The genome of the model beetle and pest Tribolium castaneum.

          Tribolium castaneum is a member of the most species-rich eukaryotic order, a powerful model organism for the study of generalized insect development, and an important pest of stored agricultural products. We describe its genome sequence here. This omnivorous beetle has evolved the ability to interact with a diverse chemical environment, as shown by large expansions in odorant and gustatory receptors, as well as P450 and other detoxification enzymes. Development in Tribolium is more representative of other insects than is Drosophila, a fact reflected in gene content and function. For example, Tribolium has retained more ancestral genes involved in cell-cell communication than Drosophila, some being expressed in the growth zone crucial for axial elongation in short-germ development. Systemic RNA interference in T. castaneum functions differently from that in Caenorhabditis elegans, but nevertheless offers similar power for the elucidation of gene function and identification of targets for selective insect control.
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            Immune pathways and defence mechanisms in honey bees Apis mellifera

            Social insects are able to mount both group-level and individual defences against pathogens. Here we focus on individual defences, by presenting a genome-wide analysis of immunity in a social insect, the honey bee Apis mellifera. We present honey bee models for each of four signalling pathways associated with immunity, identifying plausible orthologues for nearly all predicted pathway members. When compared to the sequenced Drosophila and Anopheles genomes, honey bees possess roughly one-third as many genes in 17 gene families implicated in insect immunity. We suggest that an implied reduction in immune flexibility in bees reflects either the strength of social barriers to disease, or a tendency for bees to be attacked by a limited set of highly coevolved pathogens.
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              Social immunity.

              Social insect colonies have evolved collective immune defences against parasites. These 'social immune systems' result from the cooperation of the individual group members to combat the increased risk of disease transmission that arises from sociality and group living. In this review we illustrate the pathways that parasites can take to infect a social insect colony and use these pathways as a framework to predict colony defence mechanisms and present the existing evidence. We find that the collective defences can be both prophylactic and activated on demand and consist of behavioural, physiological and organisational adaptations of the colony that prevent parasite entrance, establishment and spread. We discuss the regulation of collective immunity, which requires complex integration of information about both the parasites and the internal status of the insect colony. Our review concludes with an examination of the evolution of social immunity, which is based on the consequences of selection at both the individual and the colony level.
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                Author and article information

                Contributors
                Journal
                Front Genet
                Front Genet
                Front. Genet.
                Frontiers in Genetics
                Frontiers Media S.A.
                1664-8021
                04 March 2015
                2015
                : 6
                : 9
                Affiliations
                [1] 1Department of Evolutionary Biology and Ecology, Institute of Biology I, University of Freiburg Freiburg, Germany
                [2] 2Section for Ecology and Evolution, Department of Biology, Centre for Social Evolution, University of Copenhagen Copenhagen, Denmark
                [3] 3China National Genebank, BGI-Shenzhen Shenzhen, China
                [4] 4Centre for GeoGenetics, Natural History Museum of Denmark, University of Copenhagen Copenhagen, Denmark
                [5] 5School of Life Sciences, Arizona State University Tempe, AZ, USA
                Author notes

                Edited by: Juergen Rudolf Gadau, Arizona State University, USA

                Reviewed by: Seirian Sumner, University of Bristol, UK; Bart Pannebakker, Wageningen University, Netherlands; Michael E. Scharf, Purdue University, USA

                *Correspondence: Judith Korb, Department of Evolutionary Biology and Ecology, Institute of Biology I, University of Freiburg, Hauptstrasse 1, D-79104 Freiburg, Germany e-mail: judith.korb@ 123456biologie.uni-freiburg.de

                This article was submitted to Evolutionary and Population Genetics, a section of the journal Frontiers in Genetics.

                Article
                10.3389/fgene.2015.00009
                4348803
                25788900
                cb647683-7e24-4a68-837f-f8d61f7fcb91
                Copyright © 2015 Korb, Poulsen, Hu, Li, Boomsma, Zhang and Liebig.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 29 September 2014
                : 09 January 2015
                Page count
                Figures: 4, Tables: 5, Equations: 0, References: 101, Pages: 12, Words: 8983
                Categories
                Genetics
                Original Research Article

                Genetics
                chemical communication,genomes,immunity,social organization,social insects,symbiosis,termites,transposable elements

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