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      The prospects of selection for social genetic effects to improve welfare and productivity in livestock

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          Abstract

          Social interactions between individuals living in a group can have both positive and negative effects on welfare, productivity, and health of these individuals. Negative effects of social interactions in livestock are easier to observe than positive effects. For example, laying hens may develop feather pecking, which can cause mortality due to cannibalism, and pigs may develop tail biting or excessive aggression. Several studies have shown that social interactions affect the genetic variation in a trait. Genetic improvement of socially-affected traits, however, has proven to be difficult until relatively recently. The use of classical selection methods, like individual selection, may result in selection responses opposite to expected, because these methods neglect the effect of an individual on its group mates (social genetic effects). It has become clear that improvement of socially-affected traits requires selection methods that take into account not only the direct effect of an individual on its own phenotype but also the social genetic effects, also known as indirect genetic effects, of an individual on the phenotypes of its group mates. Here, we review the theoretical and empirical work on social genetic effects, with a focus on livestock. First, we present the theory of social genetic effects. Subsequently, we evaluate the evidence for social genetic effects in livestock and other species, by reviewing estimates of genetic parameters for direct and social genetic effects. Then we describe the results of different selection experiments. Finally, we discuss issues concerning the implementation of social genetic effects in livestock breeding programs. This review demonstrates that selection for socially-affected traits, using methods that target both the direct and social genetic effects, is a promising, but sometimes difficult to use in practice, tool to simultaneously improve production and welfare in livestock.

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          Best linear unbiased estimation and prediction under a selection model.

          Mixed linear models are assumed in most animal breeding applications. Convenient methods for computing BLUE of the estimable linear functions of the fixed elements of the model and for computing best linear unbiased predictions of the random elements of the model have been available. Most data available to animal breeders, however, do not meet the usual requirements of random sampling, the problem being that the data arise either from selection experiments or from breeders' herds which are undergoing selection. Consequently, the usual methods are likely to yield biased estimates and predictions. Methods for dealing with such data are presented in this paper.
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            Evolutionary consequences of indirect genetic effects

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              Multilevel selection 1: Quantitative genetics of inheritance and response to selection.

              Interaction among individuals is universal, both in animals and in plants, and substantially affects evolution of natural populations and responses to artificial selection in agriculture. Although quantitative genetics has successfully been applied to many traits, it does not provide a general theory accounting for interaction among individuals and selection acting on multiple levels. Consequently, current quantitative genetic theory fails to explain why some traits do not respond to selection among individuals, but respond greatly to selection among groups. Understanding the full impacts of heritable interactions on the outcomes of selection requires a quantitative genetic framework including all levels of selection and relatedness. Here we present such a framework and provide expressions for the response to selection. Results show that interaction among individuals may create substantial heritable variation, which is hidden to classical analyses. Selection acting on higher levels of organization captures this hidden variation and therefore always yields positive response, whereas individual selection may yield response in the opposite direction. Our work provides testable predictions of response to multilevel selection and reduces to classical theory in the absence of interaction. Statistical methodology provided elsewhere enables empirical application of our work to both natural and domestic populations.
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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
                11 November 2014
                2014
                : 5
                : 377
                Affiliations
                [1] 1Animal Breeding and Genomics Centre, Wageningen University Wageningen, Netherlands
                [2] 2Behavioural Ecology Group, Wageningen University Wageningen, Netherlands
                [3] 3Hendrix Genetics, Research and Technology Centre Boxmeer, Netherlands
                [4] 4Adaptation Physiology Group, Wageningen University Wageningen, Netherlands
                [5] 5TOPIGS Research Centre IPG Beuningen, Netherlands
                [6] 6Department of Animal Science, Purdue University West Lafayette, IN, USA
                [7] 7Institut de Sélection Animale Boxmeer, Netherlands
                Author notes

                Edited by: Wendy Mercedes Rauw, Instituto Nacional de Investigación y Tecnología Agraria y Alimentaria, Spain

                Reviewed by: Rodolfo Juan Carlos Cantet, Universidad de Buenos Aires, Argentina; Jörn Bennewitz, University Hohenheim, Germany

                *Correspondence: Esther D. Ellen, Animal Breeding and Genomics Centre, Wageningen University, Radix Building number 107, Droevendaalsesteeg 1, PO Box 338, 6708 PB Wageningen, Netherlands e-mail: esther.ellen@ 123456wur.nl

                This article was submitted to Livestock Genomics, a section of the journal Frontiers in Genetics.

                Article
                10.3389/fgene.2014.00377
                4227523
                25426136
                c87401ac-e306-4bd6-a199-b4ca8dce0fa1
                Copyright © 2014 Ellen, Rodenburg, Albers, Bolhuis, Camerlink, Duijvesteijn, Knol, Muir, Peeters, Reimert, Sell-Kubiak, van Arendonk, Visscher and Bijma.

                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
                : 30 June 2014
                : 13 October 2014
                Page count
                Figures: 1, Tables: 5, Equations: 12, References: 108, Pages: 14, Words: 13026
                Categories
                Genetics
                Review Article

                Genetics
                genetic selection,social genetic effects,welfare,laying hens,pigs
                Genetics
                genetic selection, social genetic effects, welfare, laying hens, pigs

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