Transcriptomic analysis of intestinal organoids, derived from pigs divergent in feed efficiency, and their response to Escherichia coli.

BACKGROUND: There is increasing interest in using intestinal organoids to study complex traits like feed efficiency (FE) and host-microbe interactions. The aim of this study was to investigate differences in the molecular phenotype of organoids derived from pigs divergent for FE as well as their responses to challenge with adherent and invasive Escherichia coli (E. coli). RESULTS: Colon and ileum tissue from low and high FE pigs was used to generate 3D organoids and two dimensional (2D) monolayers of organoid cells for E. coli challenge. Genome-wide gene expression was used to investigate molecular differences between pigs that were phenotypically divergent for FE and to study the difference in gene expression after challenge with E. coli. We showed, (1) minor differences in gene expression of colon organoids from pigs with low and high FE phenotypes, (2) that an E. coli challenge results in a strong innate immune gene response in both colon and ileum organoids, (3) that the immune response seems to be less pronounced in the colon organoids of high FE pigs and (4) a slightly stronger immune response was observed in ileum than in colon organoids. CONCLUSIONS: These findings demonstrate the potential for using organoids to gain insights into complex biological mechanisms such as FE.

Organoids: a promising new in vitro platform in livestock and veterinary research.

Organoids are self-organizing, self-renewing three-dimensional cellular structures that resemble organs in structure and function. They can be derived from adult stem cells, embryonic stem cells, or induced pluripotent stem cells. They contain most of the relevant cell types with a topology and cell-to-cell interactions resembling that of the in vivo tissue. The widespread and increasing adoption of organoid-based technologies in human biomedical research is testament to their enormous potential in basic, translational- and applied-research. In a similar fashion there appear to be ample possibilities for research applications of organoids from livestock and companion animals. Furthermore, organoids as in vitro models offer a great possibility to reduce the use of experimental animals. Here, we provide an overview of studies on organoids in livestock and companion animal species, with focus on the methods developed for organoids from a variety of tissues/organs from various animal species and on the applications in veterinary research. Current limitations, and ongoing research to address these limitations, are discussed. Further, we elaborate on a number of fields of research in animal nutrition, host-microbe interactions, animal breeding and genomics, and animal biotechnology, in which organoids may have great potential as an in vitro research tool.

Assessing the Activity of Individual Group-Housed Broilers Throughout Life using a Passive Radio Frequency Identification System-A Validation Study.

Individual data are valuable for assessing the health, welfare and performance of broilers. In particular, data on the first few days of life are needed to study the predictive value of traits recorded early in life for later life performance. However, broilers are generally kept in groups, which hampers individual identification and monitoring of animals. Sensor technologies may aid in identifying and monitoring individual animals. In this study, a passive radio frequency identification (RFID) system was implemented to record broiler activity, in combination with traditional video recordings. The two main objectives were 1) to validate the output of the RFID system by comparing it to the recorded locations on video, and 2) to assess whether the number of antennas visited per unit time could serve as a measure of activity, by comparing it to the distance recorded on video and to the distance moved as recorded using a validated ultra-wideband (UWB) tracking system. The locations recorded by the RFID system exactly matched the video in 62.5% of the cases, and in 99.2% of the cases when allowing for a deviation of one antenna grid cell. There were moderately strong Spearman rank correlations between the distance recorded with the RFID system and the distance recorded from video (rs = 0.82) and between UWB and RFID (rs = 0.70) in approximately one-hour recordings, indicating that the RFID system can adequately track relative individual broiler activity, i.e., the activity level of a broiler in comparison to its group members. As the RFID tags are small and lightweight, the RFID system is well suited for monitoring the individual activity of group-housed broilers throughout life.

The genetic architecture of socially-affected traits: a GWAS for direct and indirect genetic effects on survival time in laying hens showing cannibalism.

BACKGROUND: Cannibalism is an important welfare problem in the layer industry. Cannibalism is a social behavior where individual survival is affected by direct genetic effects (DGE) and indirect genetic effects (IGE). Previous studies analysed repeated binomial survival, instead of survival time, which improved accuracies of breeding value predictions. Our study aimed at identifying SNPs associated with DGE and IGE for survival time, and comparing results from models that analyse survival time and repeated binomial survival. METHODS: Survival data of three layer crosses (W1 * WA, W1 * WB, and W1 * WC) were used. Each individual had one survival time record and 13 monthly survival (0/1) records. Approximately 30,000 single nucleotide polymorphisms (SNPs) were included in the genome-wide association study (GWAS), using a linear mixed model for survival time, a linear mixed model and a generalized linear mixed model for repeated binomial survival (0/1). Backwards elimination was used to determine phenotypic and genetic variance explained by SNPs. RESULTS: The same quantitative trait loci were identified with all models. A SNP associated with DGE was found in cross W1 * WA, with an allele substitution effect of 22 days. This SNP explained 3% of the phenotypic variance, and 36% of the total genetic variance. Four SNPs associated with DGE were found in cross W1 * WB, with effects ranging from 16 to 35 days. These SNPs explained 1 to 6% of the phenotypic variance and 9 to 44% of the total genetic variance. Our results suggest a link of DGE and IGE for survival time in layers with the gamma-aminobutyric acid (GABA) system, since a SNP located near a gene for a GABA receptor was associated with DGE and with IGE (not significant). CONCLUSIONS: This is one of the first large studies investigating the genetic architecture of a socially-affected trait. The power to detect SNP associations was relatively low and thus we expect that many effects on DGE and IGE remained undetected. Yet, GWAS results revealed SNPs with large DGE and a link of DGE and IGE for survival time in layers with the GABAergic system, which supports existing evidence for the involvement of GABA in the development of abnormal behaviors.

Predicting direct and indirect breeding values for survival time in laying hens using repeated measures.

BACKGROUND: Minimizing bird losses is important in the commercial layer industry. Selection against mortality is challenging because heritability is low, censoring is high, and individual survival depends on social interactions among cage members. With cannibalism, mortality depends not only on an individual's own genes (direct genetic effects; DGE) but also on genes of its cage mates (indirect genetic effects; IGE). To date, studies using DGE-IGE models have focussed on survival time but their shortcomings are that censored records were considered as exact lengths of life and models assumed that IGE were continuously expressed by all cage members even after death. However, since dead animals no longer express IGE, IGE should ideally be time-dependent in the model. Neglecting censoring and timing of IGE expression may reduce accuracy of estimated breeding values (EBV). Thus, our aim was to improve prediction of breeding values for survival time in layers that present cannibalism. METHODS: We considered four DGE-IGE models to predict survival time in layers. One model was an analysis of survival time and the three others treated survival in consecutive months as a repeated binomial trait (repeated measures models). We also tested whether EBV were improved by including timing of IGE expression in the analyses. Approximate EBV accuracies were calculated by cross-validation. The models were fitted to survival data on two purebred White Leghorn layer lines W1 and WB, each having monthly survival records over 13 months. RESULTS: Including the timing of IGE expression in the DGE-IGE model reduced EBV accuracy compared to analysing survival time. EBV accuracy was higher when repeated measures models were used. However, there was no universal best model. Using repeated measures instead of analysing survival time increased EBV accuracy by 10 to 21 and 2 to 12 % for W1 and WB, respectively. We showed how EBV and variance components estimated with repeated measures models can be translated into survival time. CONCLUSIONS: Our results suggest that prediction of breeding values for survival time in laying hens can be improved using repeated measures models. This is an important result since more accurate EBV contribute to higher rates of genetic gain.

Plumage condition in laying hens: genetic parameters for direct and indirect effects in two purebred layer lines.

BACKGROUND: Feather pecking is a major welfare issue in laying hen industry that leads to mortality. Due to a ban on conventional cages in the EU and on beak trimming in some countries of the EU, feather pecking will become an even bigger problem. Its severity depends both on the victim receiving pecking and on its group mates inflicting pecking (indirect effects), which together determine plumage condition of the victim. Plumage condition may depend, therefore, on both the direct genetic effect of an individual itself and on the indirect genetic effects of its group mates. Here, we present estimated genetic parameters for direct and indirect effects on plumage condition of different body regions in two purebred layer lines, and estimates of genetic correlations between body regions. METHODS: Feather condition scores (FCS) were recorded at 40 weeks of age for neck, back, rump and belly and these four scores were added-up into a total FCS. A classical animal model and a direct-indirect effects model were used to estimate genetic parameters for FCS. In addition, a bivariate model with mortality (0/1) was used to account for mortality before recording FCS. Due to mortality during the first 23 weeks of laying, 5363 (for W1) and 5089 (for WB) FCS records were available. RESULTS: Total heritable variance for FCS ranged from 1.5% to 9.8% and from 9.8% to 53.6% when estimated respectively with the classical animal and the direct-indirect effects model. The direct-indirect effects model had a significantly higher likelihood. In both lines, 70% to 94% of the estimated total heritable variation in FCS was due to indirect effects. Using bivariate analysis of FCS and mortality did not affect estimates of genetic parameters. Genetic correlations were high between adjacent regions for FCS on neck, back, and rump but moderate to low for belly with other regions. CONCLUSION: Our results show that 70% to 94% of the heritable variation in FCS relates to indirect effects, indicating that methods of genetic selection that include indirect genetic effects offer perspectives to improve plumage condition in laying hens. This, in turn could reduce a major welfare problem.

Research Note: Effects of high barn temperature on group-level dispersion and individual activity in broiler chickens.

Heat stress in broilers is a pressing issue in the changing climate. Data on broiler behavior might be useful for early detection of heat stress and subsequent intervention, and may provide potential indicators for heat tolerance that can be used in broiler breeding programs. Here, we used bird location data collected in a previous study during which broilers were inadvertently exposed to high ambient temperatures due to a local heat wave. We examined whether broiler behavior changed with increasing ambient temperatures, focusing on group-level dispersion behavior and individual-level locomotor activity. We observed that birds moved closer together with increasing temperatures up to 9 °C above the desired level, and remained in similar proximity or moved further apart at temperatures above that threshold. The activity level decreased or remained stable with increasing temperature during most parts of the day, but increased at the end of the day. Possibly, the birds exhibited compensatory behavior (such as drinking and eating) during the periods when the barn cooled down after a hot day, but that could not be confirmed as no behavioral observations were available. The difference in activity levels between individuals accounted for 8.4% of the total variation, suggesting that activity might be an interesting indicator trait for heat tolerance in broiler chickens. Overall, the results of this study can inform the development of behavior-based 1) early-warning systems for heat stress and 2) heat tolerance indicators, although data on behaviors that are more specific to heat stress are probably required.

Automated pose estimation reveals walking characteristics associated with lameness in broilers.

Walking ability of broilers can be improved by selective breeding, but large-scale phenotypic records are required. Currently, gait of individual broilers is scored by trained experts, however, precision phenotyping tools could offer a more objective and high-throughput alternative. We studied whether specific walking characteristics determined through pose estimation are linked to gait in broilers. We filmed male broilers from behind, walking through a 3 m × 0.4 m (length × width) corridor one by one, at 3 time points during their lifetime (at 14, 21, and 33 d of age). We used a deep learning model, developed in DeepLabCut, to detect and track 8 keypoints (head, neck, left and right knees, hocks, and feet) of broilers in the recorded videos. Using the keypoints of the legs, 6 pose features were quantified during the double support phase of walking, and 1 pose feature was quantified during steps, at maximum leg lift. Gait was scored on a scale from 0 to 5 by 4 experts, using the videos recorded on d 33, and the broilers were further classified as having either good gait (mean gait score ≤2) or suboptimal gait (mean gait score >2). The relationship of pose features on d 33 with gait was analyzed using the data of 84 broilers (good gait: 57.1%, suboptimal gait: 42.9%). Birds with suboptimal gait had sharper hock joint lateral angles and lower hock-feet distance ratios during double support on d 33, on average. During steps, relative step height was lower in birds with suboptimal gait. Step height and hock-feet distance ratio showed the largest mean deviations in broilers with suboptimal gait compared to those with good gait. We demonstrate that pose estimation can be used to assess walking characteristics during a large part of the productive life of broilers, and to phenotype and monitor broiler gait. These insights can be used to understand differences in the walking patterns of lame broilers, and to build more sophisticated gait prediction models.

Early locomotor activity in broilers and the relationship with body weight gain.

Fast-growing broilers are relatively inactive and this is thought to be a result of selection for high growth rates. This reduced activity level is considered a major cause of leg weakness and associated leg health problems. Increased activity, especially early in life, is suggested to have positive effects on leg health, but the relationship between early activity and growth is unclear. A clearer understanding of the relationship between activity early in life and body weight gain could help determine how selecting on increased early activity could affect body weight gain in broilers. Here, a radio frequency identification (RFID) tracking system was implemented to record daily individual broiler activity throughout life, in 5 production rounds. As mean activity levels alone do not capture the variation in activity over time, multiple (dynamic) descriptors of activity were determined based on the individual birds' daily distances moved, focusing on the period from 0 to 15 days old. The mean, skewness, root mean square error (RMSE), autocorrelation, and entropy of (deviations in) activity were determined at the individual level, as well as the average daily gain (ADG). Relationships between activity descriptors and ADG were determined for 318 birds. Both when combining the data from the different production rounds and when taking production round and start weight into account, a negative relationship between ADG and RMSE was observed, indicating that birds that were more variable in their activity levels had a lower ADG. However, the activity descriptors, in combination with recording round and start weight, explained only a small part (8%) of the variation in ADG. Therefore, it is recommended for future research to also record other factors affecting ADG (e.g., type of feed provided and feed intake) and to model growth curves. Overall, this study suggests that increasing early activity does not necessarily negatively affect body weight gain. This could contribute to improved broiler health and welfare if selecting for increased activity has the expected positive effects on leg health.

Adaptation of Livestock to New Diets Using Feed Components without Competition with Human Edible Protein Sources-A Review of the Possibilities and Recommendations.

Livestock feed encompasses both human edible and human inedible components. Human edible feed components may become less available for livestock. Especially for proteins, this calls for action. This review focuses on using alternative protein sources in feed and protein efficiency, the expected problems, and how these problems could be solved. Breeding for higher protein efficiency leading to less use of the protein sources may be one strategy. Replacing (part of) the human edible feed components with human inedible components may be another strategy, which could be combined with breeding for livestock that can efficiently digest novel protein feed sources. The potential use of novel protein sources is discussed. We discuss the present knowledge on novel protein sources, including the consequences for animal performance and production costs, and make recommendations for the use and optimization of novel protein sources (1) to improve our knowledge on the inclusion of human inedible protein into the diet of livestock, (2) because cooperation between animal breeders and nutritionists is needed to share knowledge and combine expertise, and (3) to investigate the effect of animal-specific digestibility of protein sources for selective breeding for each protein source and for precision feeding. Nutrigenetics and nutrigenomics will be important tools.

The relationship between gait and automated recordings of individual broiler activity levels.

Gait, or walking ability, is an often-measured trait in broilers. Individual gait scores are generally determined manually, which can be time-consuming and subjective. Automated methods of scoring gait are available, but are often implemented at the group level. However, there is an interest in automated methods of scoring gait at the individual level. We hypothesized that locomotor activity could serve as a proxy for gait of individual broilers. Locomotor activity of 137 group-housed broilers from four crosses was recorded from approximately 16 to 32 days old, using an ultra-wideband tracking system. These birds were divided over four trials. Individual gait scores were determined at the end of the tracking period, on a scale from 0 to 5, with higher scores representing worse gait. Given the limited number of birds, birds were subsequently categorized as having a good gait (GG; scores 0-2) or a suboptimal gait (SG; scores 3-5). Relationships between activity and gait classification were studied to determine whether individual activity has the potential to serve as a proxy for gait. When comparing GG and SG birds using robust linear regression, SG birds showed a lower 1) activity around the start of tracking (estimate = -1.33 ± 0.56, P = 0.019), 2) activity near the end of tracking (estimate = -1.63 ± 0.38, P < 0.001), and 3) average activity (estimate = -1.12 ± 0.41, P = 0.007). When taking day of tracking, trial, cross and body weight category (heavy versus light at approximately 2 wk old) into account, a tendency was still observed for SG birds having lower activity levels within lightweight birds, but not within heavyweight birds. This study provides indications for activity differences between gait classifications. However, given that there was considerable overlap in activity levels between the gait classifications, future research implementing additional activity-related variables is required to allow a more complete distinction between birds with different gait classifications.

Genetic parameters for social effects on survival in cannibalistic layers: combining survival analysis and a linear animal model.

BACKGROUND: Mortality due to cannibalism in laying hens is a difficult trait to improve genetically, because censoring is high (animals still alive at the end of the testing period) and it may depend on both the individual itself and the behaviour of its group members, so-called associative effects (social interactions). To analyse survival data, survival analysis can be used. However, it is not possible to include associative effects in the current software for survival analysis. A solution could be to combine survival analysis and a linear animal model including associative effects. This paper presents a two-step approach (2STEP), combining survival analysis and a linear animal model including associative effects (LAM). METHODS: Data of three purebred White Leghorn layer lines from Institut de Sélection Animale B.V., a Hendrix Genetics company, were used in this study. For the statistical analysis, survival data on 16,780 hens kept in four-bird cages with intact beaks were used. Genetic parameters for direct and associative effects on survival time were estimated using 2STEP. Cross validation was used to compare 2STEP with LAM. LAM was applied directly to estimate genetic parameters for social effects on observed survival days. RESULTS: Using 2STEP, total heritable variance, including both direct and associative genetic effects, expressed as the proportion of phenotypic variance, ranged from 32% to 64%. These results were substantially larger than when using LAM. However, cross validation showed that 2STEP gave approximately the same survival curves and rank correlations as LAM. Furthermore, cross validation showed that selection based on both direct and associative genetic effects, using either 2STEP or LAM, gave the best prediction of survival time. CONCLUSION: It can be concluded that 2STEP can be used to estimate genetic parameters for direct and associative effects on survival time in laying hens. Using 2STEP increased the heritable variance in survival time. Cross validation showed that social genetic effects contribute to a large difference in survival days between two extreme groups. Genetic selection targeting both direct and associative effects is expected to reduce mortality due to cannibalism in laying hens.

Can breeders solve mortality due to feather pecking in laying hens?

Mortality due to feather pecking (FP) has large economic and welfare consequences in the commercial poultry industry, and reduces survival of birds. With FP, the survival time of a hen depends both on her own genetic ability to avoid becoming the victim of FP (direct genetic effect; DGE), and on the genetic tendency of her group mates to perform FP (indirect genetic effect; IGE). Thus, to improve survival time of laying hens, it is important to use a breeding strategy that captures both the DGE and the IGE of selection candidates. Here, we investigate the prospects for solving mortality due to FP in laying hens by genetic selection. First, we review genetic parameters for survival time. Second, we use deterministic simulation to predict response to selection for 2 multi-trait crossbred breeding programs, a traditional recurrent testing scheme (RT) and a genomic selection scheme (GS). Finally, we investigate the prospects for sustained improvement of survival time when mortality becomes low. Results show that survival time has considerable heritable variation; most estimates of the total additive genetic standard deviation are larger than 1 mo. As expected, predicted single generation response to selection in survival time with GS is substantial larger than with RT. Particularly when the correlation between survival time and other breeding goal traits is zero, the GS scheme yields substantial improvement in survival time. For example, when mortality is 35%, the genetic correlation between survival time and other traits is 0, allowing for a 10% reduction of response in other traits, and when selection takes place in both the sire line and dam line, survival time can be improved with ∼23 D in one generation, using GS. Results, however, also show a strong decrease in heritability when mortality decreases, indicating that continued improvement becomes increasingly difficult. In summary, our results show that breeders can considerably reduce mortality due to FP with limited reduction of improvement in other breeding goal traits.

Validation of an Ultra-Wideband Tracking System for Recording Individual Levels of Activity in Broilers.

Individual data on activity of broilers is valuable, as activity may serve as a proxy for multiple health, welfare and performance indicators. However, broilers are often kept in large groups, which makes it difficult to identify and monitor them individually. Sensor technologies might offer solutions. Here, an ultra-wideband (UWB) tracking system was implemented with the goal of validating this system for individual tracking of activity of group-housed broilers. The implemented approaches were (1) a comparison of distances moved as recorded by the UWB system and on video and (2) a study recording individual levels of activity of broilers and assessing group-level trends in activity over time; that could be compared to activity trends from literature. There was a moderately strong positive correlation between the UWB system and video tracking. Using the UWB system, we detected reductions in activity over time and we found that lightweight birds were on average more active than heavier birds. Both findings match with reports in literature. Overall, the UWB system appears well-suited for activity monitoring in broilers, when the settings are kept the same for all individuals. The longitudinal information on differences in activity can potentially be used as proxy for health, welfare and performance; but further research into individual patterns in activity is required.

Review of Sensor Technologies in Animal Breeding: Phenotyping Behaviors of Laying Hens to Select Against Feather Pecking.

Damaging behaviors, like feather pecking (FP), have large economic and welfare consequences in the commercial laying hen industry. Selective breeding can be used to obtain animals that are less likely to perform damaging behavior on their pen-mates. However, with the growing tendency to keep birds in large groups, identifying specific birds that are performing or receiving FP is difficult. With current developments in sensor technologies, it may now be possible to identify laying hens in large groups that show less FP behavior and select them for breeding. We propose using a combination of sensor technology and genomic methods to identify feather peckers and victims in groups. In this review, we will describe the use of "-omics" approaches to understand FP and give an overview of sensor technologies that can be used for animal monitoring, such as ultra-wideband, radio frequency identification, and computer vision. We will then discuss the identification of indicator traits from both sensor technologies and genomics approaches that can be used to select animals for breeding against damaging behavior.

Genetic improvement of traits affected by interactions among individuals: Sib selection schemes.

Livestock populations are usually kept in groups. As a consequence, social interactions among individuals affect productivity, health, and welfare. Current selection methods (individual selection), however, ignore those interactions and yield suboptimal or in some cases even negative responses. In principle, selection between groups instead of individuals offers a solution, but has rarely been adopted in practice for two reasons. First, the relationship between group selection theory and common animal breeding concepts, such as the accuracy of selection, is unclear. Second, application of group selection requires keeping selection candidates in groups, which is often undesirable in practice. This work has two objectives. First, we derive expressions for the accuracy of individual and group selection, which provides a measurement of quality for those methods. Second, we investigate the opportunity to improve traits affected by interactions by using information on relatives kept in family groups, while keeping selection candidates individually. The accuracy of selection based on relatives is shown to be an analogy of the classical expression for traits not affected by interactions. Our results show that selection based on relatives offers good opportunities for effective genetic improvement of traits affected by interactions.

Multilevel selection 2: Estimating the genetic parameters determining inheritance and response to selection.

Interactions among individuals are universal, both in animals and in plants and in natural as well as domestic populations. Understanding the consequences of these interactions for the evolution of populations by either natural or artificial selection requires knowledge of the heritable components underlying them. Here we present statistical methodology to estimate the genetic parameters determining response to multilevel selection of traits affected by interactions among individuals in general populations. We apply these methods to obtain estimates of genetic parameters for survival days in a population of layer chickens with high mortality due to pecking behavior. We find that heritable variation is threefold greater than that obtained from classical analyses, meaning that two-thirds of the full heritable variation is hidden to classical analysis due to social interactions. As a consequence, predicted responses to multilevel selection applied to this population are threefold greater than classical predictions. This work, combined with the quantitative genetic theory for response to multilevel selection presented in an accompanying article in this issue, enables the design of selection programs to effectively reduce competitive interactions in livestock and plants and the prediction of the effects of social interactions on evolution in natural populations undergoing multilevel selection.

Direct and indirect genetic effects in life-history traits of flour beetles (Tribolium castaneum).

Indirect genetic effects (IGEs) are the basis of social interactions among conspecifics, and can affect genetic variation of nonsocial and social traits. We used flour beetles (Tribolium castaneum) of two phenotypically distinguishable populations to estimate genetic (co)variances and the effect of IGEs on three life-history traits: development time (DT), growth rate (GR), and pupal body mass (BM). We found that GR was strongly affected by social environment with IGEs accounting for 18% of the heritable variation. We also discovered a sex-specific social effect: male ratio in a group significantly affected both GR and BM; that is, beetles grew larger and faster in male-biased social environments. Such sex-specific IGEs have not previously been demonstrated in a nonsocial insect. Our results show that beetles that achieve a higher BM do so via a slower GR in response to social environment. Existing models of evolution in age-structured or stage-structured populations do not account for IGEs of social cohorts. It is likely that such IGEs have played a key role in the evolution of developmental plasticity shown by Tenebrionid larvae in response to density. Our results document an important source of genetic variation for GR, often overlooked in life-history theory.

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

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.

Selection for low mortality in laying hens affects catecholamine levels in the arcopallium, a brain area involved in fear and motor regulation.

Feather pecking (FP) in laying hens may cause mortality due to cannibalism. Novel breeding methods using survival days of group-housed siblings allow for the genetic selection of laying hens with low mortality (LML: low mortality line) due to cannibalism. Previous studies have demonstrated less fear-related behavior and also less FP in LML hens compared to CL. Selection also caused changes in locomotor behavior in an open field. It is unknown, however, whether selection for low mortality affects central neurotransmitter levels. In this study, brain monoamine levels were measured in the dorsal thalamus, medial striatum, hippocampus and arcopallium of adult laying hens of both LML and CL using HPLC. Brain samples were collected after 5-min of manual restraint. The most prominent line differences were found in the arcopallium. Compared to CL, LML had lower levels of noradrenaline (NA) and 3,4-dihydroxyphenylacetic acid (DOPAC) and tended to have lower levels of dopamine (DA), homovanillic acid (HVA), and 5-hydroxyindoleacetic acid (5-HIAA). Levels of serotonin (5-HT), 5-HT- and DA-turnover in this brain area were not affected by line. LML showed less fear-related behavior during the restraint than CL. These findings show that selection for low mortality in hens leads to changes of predominantly the dopaminergic system in the chicken's arcopallium, a forebrain somatomotor area also related to fear. This suggests a relationship between catecholamine functioning in this brain area and FP and cannibalistic behavior in chickens and underpins previously found relationships between FP, fear and high activity.