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This review focuses on the role of genes, epigenetics and ontogeny in behavioural development of animals. The behavioural characteristics of an individual are determined by its genes and by its physical and social environment. Not only the individual's early life and current environment are of importance, but also the environment of previous generations. Through epigenetic processes, stress in parents and even grandparents can translate into changes in behavioural and physical characteristics of the offspring. Another influential factor for behavioural development is maternal hormones. Recent studies indicate that hormonal effects may also be closely related to epigenetic changes. Also, the environment during ontogeny has considerable impact on behavioural development: in both mice and laying hens, high quality maternal care resulted in animals that were less fearful. In laying hens maternal care also led to a reduction in cannibalistic pecking. Genetic selection and selection experiments will also play a key role in breeding animals for the housing systems of the future. To optimize behavioural development of farm animals and to minimize risks of damaging behaviour, integral approaches are needed that combine selection of the optimal genotype with provision of a favourable environment for parents and offspring, both during ontogeny and later life.

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.

Purebred laying hen lines of White Leghorn (WL) origin have been found to be more flighty and to show more feather pecking than lines of Rhode Island Red (RIR) origin. It has been found, however, that when RIR birds were housed together with WL birds, RIR birds became more flighty and those mixed groups developed more feather damage than pure-line cage-housed groups. It is unknown, however, whether this effect of social environment is accompanied by changes in stress-related behavior and neurophysiological activity, which are assumed to be associated with increased feather damage. The objective of this study was therefore to investigate the effects of genetic origin (WL or RIR) and social environment (mixed or pure groups) on behavioral response to manual restraint and monoamine functioning. Monoamine functioning was measured by brain serotonin (5-HT) and dopamine turnover. Furthermore, correlations between 5-HT turnover in the brain and peripheral measures of 5-HT in the blood were calculated. Experimental birds, housed either with other birds from the same genetic origin (pure groups) or with both RIR and WL birds (mixed groups) from hatching onward, were subjected to a manual restraint test at 47 wk of age. The WL birds struggled less during restraint and had higher dopamine and 5-HT turnover levels after restraint than did RIR birds. The WL birds also showed higher levels of platelet 5-HT uptake than did RIR birds. No effects of social environment were found. Blood and brain 5-HT measures were found to be correlated, with correlations ranging from 0.34 to 0.57, which seems to offer opportunities for less invasive peripheral indicators of 5-HT activity. In conclusion, genetic origin, but not social environment, affected the behavioral response to manual restraint and monoamine functioning in laying hens.

International audience; Under current practices, broiler parent stock (broiler breeders) encounter several welfare problems, such as feed restriction and injury during mating. Intensive selection for production traits, especially growth rate, is associated with increased nutritious requirement and thus feed consumption, but also reproductive dysfunctions and decreased sexual activity in broiler breeders. A resulting serious welfare problem is the subsequent severe feed restriction which is applied during rearing, in order to prevent health problems and to achieve better egg production. This severe feed restriction has negative effects on bird welfare as it causes chronic stress resulting from hunger. Recent research has focused on management practices to alleviate the negative effect of this feed restriction. Using both a more fibrous feed and an appetite suppressant seems most successful. There are welfare concerns involved in the mating of broiler breeders. In many countries mutilations (beak trimming, despurring and toe clipping) are carried out in male broiler breeders as a standard procedure to prevent excessive damage to the hens during mating. It has been reported that broiler breeder males show low libido but, at the same time, may be aggressive and show rough behaviour during mating, which can cause severe injuries to the hens, although some injuries may be caused by over-mating hens. Current studies aim to develop management practices to reduce hen injury due to over-mating and improve the sexual behaviour in both males and females. In the last part of the review, an examination of how breeding companies are taking welfare into consideration in their commercial selection scheme is made. In practice, although for example reduced mortality and some health criteria are included in selection programs and some alternative breeds as used, other welfare issues remain, such as feed restriction during rearing.

An experiment was performed to investigate the effect of animal versus vegetable protein sources in the diet on the development of behavior in laying hens. A diet containing protein sources of only vegetable origin was compared with four diets, each containing one of four processed animal proteins (PAPs). Two PAPs (Daka-58 and Sonac-60) were classified as meat meals, whereas the remaining ones (Daka-40 and Sonac-50) were classified as meat and bone meals. The impact of a control diet (without PAP) versus four PAP diets (50 g/kg CP of animal origin) on behavior was determined. All diets were isocaloric (AME n = 11.8 MJ/kg) and isonitrogenous (dig. lysine = 6.8 g/kg). Hens were housed in 40 floor pens (12 hens/pen, 8 pens/treatment) from 20 to 40 weeks of age. Supplementation of PAPs did not generally reduce feather pecking behavior. Nevertheless, Daka-40 and Sonac-50 fed hens showed a delay in the development of feather damage and, simultaneously, an increase in litter condition, foraging and walking behavior, and floor pecks compared to hens fed Sonac-60. These shifts seemed to be partly related with the intake of digestible glycine, available phosphorus, calcium, potassium, and sodium.

Welfare of animals can be defined as the kind of feelings the environmental conditions bring about in the animals. These feelings depend on the needs of the animals and their degree of satisfaction. Needs of animals, and so their welfare, are partly genetically determined. Therefore, welfare can be changed by breeding. The aim of this study was to investigate how welfare of pigs under modern intensive farm conditions can be improved by genetic selection, with emphasis on the precise definition of the breeding goal and determination of the animal characteristics on which selection can be based in practice. The existing thermoregulation model was used to develop a conceptual framework that describes welfare of growing pigs and production sows with respect to each of their needs as a curvilinear function of the respective environmental conditions. The framework assumes that welfare in terms of feelings is reflected by the physiological and behavioural mechanisms the pig has to activate in order to cope with the various environmental conditions it encounters. Based on those physiological and behavioural responses to changing conditions, five welfare zones can be distinguished for each need. Breeding goals for welfare were defined in terms of the transition points between these welfare zones, such that future pigs would better cope with unfavourable or unfamiliar farming conditions, therewith quickening the domestication process, to some extent. However, as long as genetic parameters for these transition points are not available, more common welfare-related characteristics like temperament, stress resistance and robustness can be included in the breeding goal, as an alternative. For selection among potential breeding candidates, transition points between welfare zones can be determined in sib tests, thereby also collecting the data for estimating genetic parameters. As a cheaper alternative, breeding candidates could be tested under hard conditions and selected on their coping success. In addition, various behavioural tests and operant conditioning tests ( to test a pig's motivation to change its actual environment) can be carried out. Under common conditions on the farm, problems associated with coping (like incidences of diseases, injuries, and stereotypies) and/or other relevant traits ( e. g. saliva cortisol levels, longevity and even production traits) should be recorded routinely and used as selection index information. Selection for improved welfare should lead to more tolerant pigs that are better able to cope with possible unfavourable farm conditions by a more efficient use of the adaptation mechanisms they already possess. It should, however, not result in lowering husbandry standards. More research is needed to assess genetic correlations among various welfare aspects and with production traits to prevent undesired side effects in future populations of pigs.

BACKGROUND: Feather pecking and cannibalism are major concerns in poultry farming, both in terms of animal welfare and farm economics. Genetic selection and introduction of (aspects of) maternal care have been suggested as potential interventions to reduce feather pecking in laying hens. Altered brain development has been proposed to reflect welfare states in animals, and can provide more insight into the underlying processes involved in feather pecking. Both vasotocin (the avian homologue of vasopressin) and dopaminergic neural circuitry have roles in control of social behaviors as well as in the stress response, and may be linked to feather pecking. Thus, the hypothalamus of adult laying hens selected for low early mortality (LML), which show low feather pecking, was examined and compared with a control line of adult laying hens selected for production characteristics only (CL). The effect of foster hen rearing on the two genetic lines and their hypothalamic morphology was also investigated. RESULTS: We demonstrated an increase in the number of neurons positive for the rate-limiting enzyme in dopamine production, tyrosine hydroxylase, in the periventricular area of the hypothalamus in the LML hens compared to CL hens. Hen-reared chicks showed more vasotocin -positive neurons in the medial pre-optic area compared to the hens raised without a hen. No correlations were found between behavior in an open field at 5-6 weeks of age, and the histology of the same hens at adulthood. CONCLUSION: The hypothalamic dopaminergic and vasotinergic systems are altered in hens following genetic selection or maternal care, indicating a potential role for these systems in feather pecking. Keywords: Hypothalamus, Vasotocin, Vasopressin, Tyrosine hydroxylase, Dopamine, Welfare, Laying hen

International audience; 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.MethodsFeather 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.ResultsTotal 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.ConclusionOur 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.

The detection of quantitative trait loci (QTL) of behavioural traits has mainly been focussed on mouse and rat. With the rapid development of molecular genetics and the statistical tools, QTL mapping for behavioural traits in farm animals is developing. In chicken, a total of 30 QTL involved in pecking-related traits, open-field behaviour, tonic immobility, response to novel objects, and response to a restraint test were detected in different studies. In the search for a useful early predictor for feather pecking (FP) behaviour in adult laying hens, the following was found: FP in young animals is not a predictor for FP in adult animals, however, open-field behaviour in young animals is genetically correlated with FP in adult hens. Before the implementation of FP behaviour or open-field behaviour in breeding programmes, it is essential to know more about the correlation between these behavioural traits and also their relationship with production traits. Nevertheless, with the QTL for behavioural traits and the chicken genome sequence in progress, a better understanding of the underlying genetic mechanisms of behavioural traits will be feasible.

Line differences for open-field behavior in chickens have been observed, and it has been shown that this behavior has a genetic component. The aim of this study was to detect quantitative trait loci (QTL) involved in open-field behavior. For this purpose, open-field behavior was studied at 5 and 29 weeks of age in F(2) hens coming from an intercross between two commercial White Leghorn laying lines selected for egg production traits. Latencies, durations, and frequencies of general activity (sitting, standing, walking, and stepping), defecation, and vocalizations were recorded individually for each bird, and a factor score was calculated. All animals (F(0), F(1), and F(2)) were screened with 180 microsatellite markers. Regression interval mapping was applied using both a paternal half-sib analysis and a line-cross analysis method. For general activity at 5 weeks of age, a significant QTL was detected on GGA4 and a suggestive QTL on GGA2 under the line-cross model. For general activity at 29 weeks of age, a significant QTL was detected on GGA4 and two suggestive QTLs were detected on GGA1 and on GGA10, respectively, also using the line-cross analysis. The QTL on GGA4 at 5 weeks of age did not overlap with the QTL on GGA4 at 29 weeks of age. The current study indicated that open-field behavior in young chickens was regulated by QTL that differ from the QTL for open-field behavior in adult chickens.

Welfare of animals can be defined as the kind of feelings the environmental conditions bring about in the animals. These feelings depend on the needs of the animals and their degree of satisfaction. Needs of animals, and so their welfare, are partly genetically determined. Therefore, welfare can be changed by breeding. The aim of this study was to investigate how welfare of pigs under modern intensive farm conditions can be improved by genetic selection, with emphasis on the precise definition of the breeding goal and determination of the animal characteristics on which selection can be based in practice.The existing thermoregulation model was used to develop a conceptual framework that describes welfare of growing pigs and production sows with respect to each of their needs as a curvilinear function of the respective environmental conditions. The framework assumes that welfare in terms of feelings is reflected by the physiological and behavioural mechanisms the pig has to activate in order to cope with the various environmental conditions it encounters. Based on those physiological and behavioural responses to changing conditions, five welfare zones can be distinguished for each need. Breeding goals for welfare were defined in terms of the transition points between these welfare zones, such that future pigs would better cope with unfavourable or unfamiliar farming conditions, therewith quickening the domestication process, to some extent. However, as long as genetic parameters for these transition points are not available, more common welfare-related characteristics like temperament, stress resistance and robustness can be included in the breeding goal, as an alternative.For selection among potential breeding candidates, transition points between welfare zones can be determined in sib tests, thereby also collecting the data for estimating genetic parameters. As a cheaper alternative, breeding candidates could be tested under hard conditions and selected on their coping success. In addition, various behavioural tests and operant conditioning tests (to test a pig's motivation to change its actual environment) can be carried out. Under common conditions on the farm, problems associated with coping (like incidences of diseases, injuries, and stereotypies) and/or other relevant traits (e.g. saliva cortisol levels, longevity and even production traits) should be recorded routinely and used as selection index information. Selection for improved welfare should lead to more tolerant pigs that are better able to cope with possible unfavourable farm conditions by a more efficient use of the adaptation mechanisms they already possess. It should, however, not result in lowering husbandry standards. More research is needed to assess genetic correlations among various welfare aspects and with production traits to prevent undesired side effects in future populations of pigs.