Sexual dimorphism in immune function influences disease susceptibility, yet sex-specific immune characteristics in chickens remain poorly understood, particularly across genetically divergent lines. This study investigated sex- and line-dependent differences in the peripheral and lymphatic immune system in lines selected for high and low feather pecking. Immune cell subsets were quantified in blood, spleen and cecal tonsils, and functional immune parameters were analyzed by measuring plasma antibody concentration and mitogen-induced lymphocyte proliferation capacity. Distinct immunological differences were observed between roosters and hens, as well as between lines, with several effects being tissue- and sex-specific. The higher overall numbers of immune cells observed in the spleen of roosters were primarily attributable to their larger organ mass, whereas in the cecal tonsils this was mainly due to a higher cell density. Roosters had fewer peripheral B cells, lower antibody concentrations and B lymphocyte proliferation capacity, along with fewer CD4+ cells and a lower CD4+/CD8α+ ratio. However, they had more γδ T cells compared to hens, indicating a greater reliance on cytotoxic or innate immune mechanisms rather than on adaptive, B cell mediated immunity. Line-specific immune differences were also observed, with the number of CD4+ cells in the cecal tonsils being lower exclusively in high feather pecking hens, supporting the hypothesis that feather pecking might be an immune-related behavior, potentially based on immune imbalances in the gut. In conclusion, immune profiles are strongly influenced by both sex and genetic background. Our results further emphasize the importance of considering sex when analyzing immune responses, as well as in vaccination strategies, disease management, and breeding programs.

Feather pecking (FP) is a serious behavioral disorder in laying hens, leading to feather damage, skin lesions, and often resulting in cannibalism. The mechanisms underlying FP are not clear yet, but recently the role of the immune system as a cause has been discussed. In humans, the interrelation between personality traits and the immune system is well-documented, with impulsivity and hyperactivity linked to distinct alterations in blood immune cell numbers and to elevated levels of pro-inflammatory cytokines. Similarly, FP in hens is associated with impulsivity and hyperactivity, suggesting a possible connection between FP and immune cell alterations. In this study numbers of leukocyte subsets in blood, spleen and cecal tonsils, along with mitogen-induced lymphocyte proliferative response and antibody concentrations across hens selectively bred for high (HFP) and low (LFP) feather pecking behavior were analyzed. Results showed that divergent selection altered FP behavior, with HFP hens showing about 10 times more pecking behavior than hens of the LFP line. HFP hens had lower numbers of T helper cells, CD4+ CD25high as well as B cells compared to LFP hens. Furthermore, HFP hens demonstrated a stronger proliferation of T cells when stimulated with ConA, while showed a weaker response in T cell-dependent B cell proliferation when stimulated with PWM, compared to LFP hens. Antibody plasma concentrations were similar between both lines. These findings highlight substantial immunological differences between HFP and LFP hens, especially in T cell immunity, and support the hypothesis that FP may be an immune-related behavioral response.

Various aspects of activity, such as spontaneous activity, explorative activity, activity in open-field tests, and hyperactivity syndrome have been explored as causal factors of feather pecking in laying hens, with no clear results. In all previous studies, mean values of activity over different time intervals were used as criteria. Incidental observation of alternated oviposition time in lines selected for high (HFP) and low feather pecking (LFP), supported by a recent study which showed differentially expressed genes related to the circadian clock in the same lines, led to the hypothesis that feather pecking may be related to a disturbed diurnal activity rhythm. Hence activity recordings of a previous generation of these lines have been reanalyzed. Data sets of a total of 682 pullets of 3 subsequent hatches of HFP, LFP, and an unselected control line (CONTR) were used. Locomotor activity was recorded in pullets housed in groups of mixed lines in a deep litter pen on 7 consecutive 13-h light phases, using a radio-frequency identification antenna system. The number of approaches to the antenna system was recorded as a measure of locomotor activity and analyzed using a generalized linear mixed model including hatch, line, time of day and the interactions of hatch × time of day and line × time of day as fixed effects. Significant effects were found for time and the interaction line × time of day but not for line. All lines showed a bimodal pattern of diurnal activity. The peak activity of the HFP in the morning was lower than that of the LFP and CONTR. In the afternoon peak all lines differed with the highest mean in the LFP followed by CONTR and HFP. The present results provide support for the hypothesis that a disturbed circadian clock plays a role in the development of feather pecking.

Different breeds of domestic and junglefowl differ in foraging strategies indicating that domestication resulted in modified energy saving behavioral strategies. In the present study we investigated foraging strategies and foraging-related behavior in 4 lines of laying hens differing in phylogenetic origin and laying performance to analyze a possible relationship between foraging and the level of egg production. High performing brown and white pure bred lines were compared with their low performing brown and white counterparts. To control possible effects on behavior other than genetic effects, all hens were reared and kept in an identical environment. A total of 72 hens from each line were kept in 6 compartments with 12 hens per compartment, respectively. Observations were done for 3 times during one laying period. Foraging strategy was tested by a contrafreeloading (CFL) paradigm. CFL describes a behavior in which animals prefer food that requires effort to obtain, although at the same time food is freely available. The hens were offered a commercial standard diet in one trough and a mixture of wood shavings and commercial standard diet in another trough. The behavior of hens was video recorded and the activity level of individual hens in the litter area was recorded by an antenna-transponder system. The high performing layers showed less CFL and foraging-related behavior compared with their low performing counterparts in both the white and brown layers. Despite differences in CFL, all hens showed a preference for the commercial standard diet compared to the mixture of wood-shavings. Our results show an association between foraging strategy and level of egg production. This suggests that a high level of egg production is accompanied by behaviors enabling the hens to satisfy their higher energy demand more efficiently. Saving energy by reduced activity probably allows them to reallocate energy into reproduction, that is, laying performance.

Killing male chicks of layer lines (MLL) is being debated under animal welfare and ethical aspects. This procedure has been banned in Germany as of January 2022 and other European countries may also consider a ban. It is known that growth and feed conversion rate in layer breeds are inferior to the performance of conventional broilers (CBR). The environmental impact of this production system has not been assessed so far. The emissions of greenhouse gases (CO(2)eq), land use, water use, acidification (SO(2)eq), and eutrophication (PO(4)eq) of MLL (Lohmann Classic, brown layer line) and CBR (Ross 308) are compared under simulated practical conditions. CBR are grown on deep litter for 32 days and fed a two-phases feeding program. Stocking density was 39 kg per m(2). MLL were grown for 90 days on deep litter using a three-phase feeding programme. Stocking density was 35 kg per m(2). CO(2)eq were calculated on the basis of CO2 emissions from respiration and litter, CO(2)eq from N2O and CH4 emissions, CO(2)eq from feed (crop production, transport and processing) and CO(2)eq from electricity and gas consumption. Based on kg slaughter yield (SY) as functional unit MLL produced more CO(2)eq (9.7 vs. 3.0 kg), used more land (17,1 vs. 4,8 m(2)) and water (196 vs 92 l) and showed a higher impact on acidification (88.6 vs. 22.7 kg SO(2)eq) and eutrophication potential (23.6 vs. 7.6 kg PO(4)eq) than CBR. The environmental effects of raising MLL for meat production are in contrast to the attempts to reduce emissions from livestock production.

Background: Feather pecking is a well-known problem in layer flocks that causes animal welfare restrictions and contributes to economic losses. Birds' gut microbiota has been linked to feather pecking. This study aims to characterize the microbial communities of two laying hen lines divergently selected for high (HFP) and low (LFP) feather pecking and investigates if the microbiota is associated with feather pecking or agonistic behavior. Methods: Besides phenotyping for the behavioral traits, microbial communities from the digesta and mucosa of the ileum and caeca were investigated using target amplicon sequencing and functional predictions. Microbiability was estimated with a microbial mixed linear model. Results: Ileum digesta showed an increase in the abundance of the genus Lactobacillus in LFP, while Escherichia was abundant in HFP hens. In the caeca digesta and mucosa of the LFP line were more abundant Faecalibacterium and Blautia. Tryptophan metabolism and lysine degradation were higher in both digesta and mucosa of the HFP hens. Linear models revealed that the two lines differ significantly in all behavior traits. Microbiabilities were close to zero and not significant in both lines and for all traits. Conclusions: Trait variation was not affected by the gut microbial composition in both selection lines.

In poultry, aggressive pecking and threatening are normal agonistic behavior patterns which serve to establish a social hierarchy. As agonistic behavior is a stressor for animals, its excessive occurrence is undesired in layer flocks. Feather pecking is a longstanding serious problem in layer flocks and its relationship to agonistic behavior is still not clear. Therefore, phenotypic and genomic analyses of the agonistic and feather pecking behavior of two laying hen lines divergently selected for high and low feather pecking were conducted. The hens were phenotyped for the active traits aggressive pecks delivered (APD)1, threats delivered (TD)2, feather pecks delivered (FPD)3 and the passive traits aggressive pecks received (APR)4, threats received (TR)5 and feather pecks received (FPR)6. Indices were built by subtracting the passive traits from the respective active traits to obtain the aggression index, the threat index, and the feather pecking index. As all three behavior patterns in their excessive manifestations are undesired, the index-traits Activity and Passivity were also defined by combining each the active and passive traits. The results showed that FPD is significantly positive correlated with APD and TD in both lines, but with higher coefficients in the high feather pecking line. The average amount of FPR in both lines is nearly the same and no correlation was found between FPR and FPD, APD or TD in any of the lines. The active traits and the feather pecking index showed medium heritabilities, whereas the heritability was negligibly small for the other traits. GWAS revealed four nominal significant (p <= 5*10(-5)) SNPs for APD on chromosome 6, the same four and three additional SNPs on chromosome 8 for Activity and three SNPs on chromosome 1 for the feather pecking index. It is concluded, that selection on high feather pecking leads to an increase of agonistic behavior. The correlation probably depends on the phase of establishing the social hierarchy in which the hens in a newly formed group are at the time of observation, and might disappear, after a stable ranking is established. The reception of feather pecking is similar in both lines. GWAS revealed that TD, APD, Activity and the feather pecking index seem to be typical quantitative traits with associated regions for the latter three which have slightly greater effects on these traits than other regions in the genome.

Feather pecking (FP) is a serious economic and welfare problem in the domestic fowl. It has recently been shown that the distribution of FP bouts within groups is heterogeneous and contains a sub-population of extreme feather peckers (EFP). The present study proposed a novel model to detect EFP hens. A mixture of two negative binomial distributions was fitted to FP data of a F2 cross of about 960 hens, and, based on the results, a calculation of the posterior probability for each hen belonging to the EFP subgroup (pEFP) was done. The fit of the mixture distribution revealed that the EFP subgroup made up a proportion of one third of the F2 cross. The EFP birds came more frequently into pecking mood and showed higher pecking intensities compared to the remaining birds. Tonic immobility and emerge box tests were conducted at juvenile and adult age of the hens to relate fearfulness to EFP. After dichotomization, all traits were analyzed in a multivariate threshold model and a genomewide association study was performed. The new trait pEFP has a medium heritability of 0.35 and is positively correlated with the fear traits. Breeding for this new trait could be an interesting option to reduce the proportion of extreme feather peckers. An index of fear related traits might serve as a proxy to breed indirectly for pEFP. GWAS revealed that all traits are typical quantitative traits with many genes and small effects contributing to the genetic variance.

Background Feather pecking (FP) in laying hens reduces animal welfare and leads to economic losses for the layer industry. FP is considered a heritable condition that is influenced by dysregulation of neurotransmitter homeostasis, the gut microbiome, and the immune system. To identify genes and biological pathways responsible for FP behavior we compared the brain transcriptomes of 48 hens divergently selected for FP. In addition, we tested if high feather peckers (HFP) and low feather peckers (LFP) respond differently to light since light has been shown to trigger FP behavior. Results Of approximately 48 million reads/sample an average of 98.4% were mapped to the chicken genome (GRCg6a). We found 13,070 expressed genes in the analyzed brains of which 423 showed differential expression between HFP and LFP. Genes of uncertain function and non-coding RNAs were overrepresented among those transcripts. Functional analyses revealed the involvement of cholinergic signaling, postsynaptic activity, membrane channels, and the immune system. After the light stimulus, 28 genes were found to be differentially expressed. These included an interaction cluster of core components of the circadian clock. However, differences in the response to light between HFP and LFP were not detectable. Conclusions Genes involved in cholinergic signaling, channel activity, synaptic transmission, and immune response were found to be involved in FP behavior. We propose a model in which the gut microbiota modulates the immune system, which in turn affects cholinergic signaling. This might have an influence on monoamine signaling with possible involvement of GABA or glutamate signaling.

Feather pecking (FP) is a longstanding serious problem in commercial flocks of laying hens. It is a highly polygenic trait and the genetic background is still not completely understood. In order to find genomic regions influencing FP, selection signatures between laying hen lines divergently selected for high and low feather pecking were mapped using the intra-population iHS and the inter-population FST approach. In addition, the existence of an extreme subgroup of FP hens (EFP) across both selected lines has been demonstrated by fitting a mixture of negative binomial distributions to the data and calculating the posterior probability of belonging to the extreme subgroup (pEFP) for each hen. A genomewide association study (GWAS) was performed for the traits pEFP and FP delivered (FPD) with a subsequent post GWAS analysis. Mapping of selection signatures revealed no clear regions under selection. GWAS revealed a region on Chromosome 1, where the existence of a QTL influencing FP is likely. The candidate genes found in this region are a part of the GABAergic system, which has already been linked to FP in previous studies. Despite the polygenic nature of FP, selection on these candidate genes may reduce FP.

BACKGROUND:Feather pecking (FP) is damaging behavior in laying hens leading to global economic losses in the layer industry and massive impairments of animal welfare. The objective of the study was to discover genetic variants and affected genes that lead to FP behavior. To achieve that we imputed low-density genotypes from two different populations of layers divergently selected for FP to sequence level by performing whole genome sequencing on founder and half-sib individuals. In order to decipher the genetic structure of FP, genome wide association studies and meta-analyses of two resource populations were carried out by focusing on the traits 'feather pecks delivered' (FPD) and the 'posterior probability of a hen to belong to the extreme feather pecking subgroup' (pEFP).RESULTS:In this meta-analysis, we discovered numerous genes that are affected by polymorphisms significantly associated with the trait FPD. Among them SPATS2L, ZEB2, KCHN8, and MRPL13 which have been previously connected to psychiatric disorders with the latter two being responsive to nicotine treatment. Gene set enrichment analysis revealed that phosphatidylinositol signaling is affected by genes identified in the GWAS and that the Golgi apparatus as well as brain structure may be involved in the development of a FP phenotype. Further, we were able to validate a previously discovered QTL for the trait pEFP on GGA1, which contains variants affecting NIPA1, KIAA1211L, AFF3, and TSGA10.CONCLUSIONS:We provide evidence for the involvement of numerous genes in the propensity to exhibit FP behavior that could aid in the selection against this unwanted trait. Furthermore, we identified variants that are involved in phosphatidylinositol signaling, Golgi metabolism and cell structure and therefore propose changes in brain structure to be an influential factor in FP, as already described in human neuropsychiatric disorders.

During their lifespan, chickens are confronted with a wide range of acute and chronic stressors in their housing environment that may threaten their welfare and health by modulating the immune system. Especially chronic stressful conditions can exceed the individual’s allostatic load, with negative consequences for immunity. A fully functional immune system is mandatory for health and welfare and, consequently, also for high productivity and safe animal products. This review provides a comprehensive overview of the impact of housing form, light regime as well as aerial ammonia and hydrogen sulfide concentrations on the immune system in chickens. Certain housing conditions are clearly associated with immunological alterations which potentially impair the success of vaccinations or affect disease susceptibility. Such poor conditions counteract sustainable poultry production. This review also outlines current knowledge gaps and provides recommendations for future research.

Animal welfare has become an important issue in poultry production. Concern about poultry welfare has mainly been expressed in industrialised countries. Since trade of poultry products is highly internationalised, welfare aspects have to be considered by all countries involved. This paper reviews the changes in the attitude to animal welfare in Western societies and the related development of regulations and standards, the impact of high welfare standards on production costs and on international trade of poultry products and finally, the influence of different stakeholders on poultry production and marketing. From the 1960s onwards, animal welfare activities have been focused on farm animals kept under ‘industrial’ conditions, such as caged laying hens. Consequently, the management conditions for laying hens in Europe have been regulated in detail by national laws and EU-Directives. Meanwhile, conventional cages have been banned in the EU and welfare activities are now directed towards other issues, such as beak-trimming and killing day-old chicks of layer lines. All measures which are considered to improve the welfare of animals increase cost of production. Hence, differences in national welfare regulations are expected to relocate poultry production to countries with low welfare standards. There is a tendency that important retailers and food chains use welfare as a marketing argument and establish high price premium labels. Standards which are established and controlled by stakeholders of the poultry market are independent of national welfare legislation. This will lead to harmonisation of welfare standards on an international level. Most welfare labels have been developed by retailers in cooperation with welfare-oriented NGOs. There is a new trend in Germany where retailers and farmers organisations develop welfare schemes and poultry producers are paid an extra allowance for welfare-friendly production. This system ensures that poultry farmers are reimbursed for the welfare-related costs.

Feather pecking is a serious damaging behaviour in the domestic fowl. Despite intensive research, the causes of this behaviour are not fully understood. Foraging and fear are considered as important causal motivations for feather pecking in the Domestic Fowl. There exist however, contrasting results which challenge the general validity of the hypotheses. It was the aim of the present study to unravel the relationships between severe feather pecking (FPD; bouts of severe feather pecks delivered), foraging (FOR, sum of walking and litter pecking), and open-field activity (OFA; number of steps) as criterion for fear, using observations of 862 birds of a F2-cross of lines selected for high or low levels of feather pecking. Heritability, phenotypic- and genetic correlations were estimated using standard multitrait analysis. Putative causal relationships between the criteria were determined using structural equation models. It was hypothesised that both OFA and FOR influence FPD, and that OFA influences FOR. Heritability for FPD and OFA was 0.19 and 0.21 respectively. The heritability for FOR was zero. Consequently, it was not possible to estimate genetic correlation between FOR and other traits. Genetic correlation between OFA and FPD as well as all phenotypic correlations among traits were very low. Recursive effects among the traits were also low and varied between 0.01 and 0.004. The results of the F2-cross used in the present study do not support the hypotheses that FOR and OPA have a causal influence on FPD.

Background Feather pecking and aggressive pecking in laying hens are serious economic and welfare issues. In spite of extensive research on feather pecking during the last decades, the motivation for this behavior is still not clear. A small to moderate heritability has frequently been reported for these traits. Recently, we identified several single-nucleotide polymorphisms (SNPs) associated with feather pecking by mapping selection signatures in two divergent feather pecking lines. Here, we performed a genome-wide association analysis (GWAS) for feather pecking and aggressive pecking behavior, then combined the results with those from the recent selection signature experiment, and linked them to those obtained from a differential gene expression study. Methods A large F2 cross of 960 F2 hens was generated using the divergent lines as founders. Hens were phenotyped for feather pecks delivered (FPD), aggressive pecks delivered (APD), and aggressive pecks received (APR). Individuals were genotyped with the Illumina 60K chicken Infinium iSelect chip. After data filtering, 29,376 SNPs remained for analyses. Single-marker GWAS was performed using a Poisson model. The results were combined with those from the selection signature experiment using Fisher’s combined probability test. Results Numerous significant SNPs were identified for all traits but with low false discovery rates. Nearly all significant SNPs were located in clusters that spanned a maximum of 3 Mb and included at least two significant SNPs. For FPD, four clusters were identified, which increased to 13 based on the meta-analysis (FPD meta ). Seven clusters were identified for APD and three for APR. Eight genes (of the 750 investigated genes located in the FPD meta clusters) were significantly differentially-expressed in the brain of hens from both lines. One gene, SLC12A9 , and the positional candidate gene for APD, GNG2 , may be linked to the monomanine signaling pathway, which is involved in feather pecking and aggressive behavior. Conclusions Combining the results from the GWAS with those of the selection signature experiment substantially increased the statistical power. The behavioral traits were controlled by many genes with small effects and no single SNP had effects large enough to justify its use in marker-assisted selection.

An important indicator of the health and behavior of laying hens is their plumage condition. Various scoring systems are used, and various risk factors for feather damage have been described. Often, a summarized score of different body parts is used to describe the overall condition of the plumage of a bird. However, it has not yet been assessed whether such a whole body plumage score is a suitable outcome variable when analyzing the risk factors for plumage deterioration. Data collected within a German project on farms keeping laying hens in aviaries were analyzed to investigate whether and the extent to which information is lost when summarizing the scores of the separate body parts. Two models were fitted using multiblock redundancy analysis, in which the first model included the whole body score as one outcome variable, while the second model included the scores of the individual body parts as multiple outcome variables. Although basically similar influences could be discovered with both models, the investigation of the individual body parts allowed for consideration of the influences on each body part separately and for the identification of additional influences. Furthermore, ambivalent influences (a factor differently associated with 2 different outcomes) could be detected with this approach, and possible dilutive effects were avoided. We conclude that influences might be underestimated or even missed when modeling their explanatory power for an overall score only. Therefore, multivariate methods that allow for the consideration of individual body parts are an interesting option when investigating influences on plumage condition.

Feather pecking is a serious economic and welfare problem in laying hens. Feather damage occurs mainly through severe feather pecking (SFP). Selection experiments have proved that this behavior is heritable and lines have been divergently selected for high (HFP) and low feather pecking (LFP). The number of bouts of SFP per hen follows a Poisson distribution with a maximum nearby 0. A few studies indicate that the distribution within flocks is not homogenous but contains sub-groups of birds showing extremely high levels of feather pecking (EFP). It was the aim of the current study to re-analyze data on SFP of lines selected for HFP/LFP and their F2 cross so as to uncover hidden sub-populations of EFP birds. Data of seven selection generations of HFP and LFP selection lines as well as their F2 cross have been used. We fitted a two-component mixture of Poisson distributions in order to separate the sub-group of EFP from the remaining birds. HFP and LFP lines differed mainly in mean bouts per bird. The proportion of EFP was only marginal in the LFP as compared with the HFP and the F2 population. Selection for LFP did not result in total elimination of EFP. The presence of even small proportions of EFP may play an important role in initiating outbreaks of feather pecking in large flocks. Further studies on feather pecking should pay special attention to the occurrence of EFP sub-groups.

The objective of this research was to analyze the relationship between feather pecking (FP) and feather eating (FE) as well as general locomotor activity (GLA) using structural equation models, which allow that one trait can be treated as an explanatory variable of another trait. This provides an opportunity to infer putative causal links among the traits. For the analysis, 897 F2-hens set up from 2 lines divergently selected for high and low FP were available. The FP observations were Box-Cox transformed, and FE and GLA observations were log and square root transformed, respectively. The estimated heritabilities of FE, GLA, and FP were 0.36, 0.29, and 0.20, respectively. The genetic correlation between FP and FE (GLA) was 0.17 (0.04). A high genetic correlation of 0.47 was estimated between FE and GLA. The recursive effect from FE to FP was [Formula: see text], and from GLA to FP [Formula: see text] These results imply that an increase of FE leads to an increased FP behavior and that an increase in GLA results in a higher FP value. Furthermore, the study showed that the genetic correlation among the traits is mainly caused by indirect effects.

Feather pecking is a well known problem in flocks of laying hens. It is partially controlled by genetics. Fear is frequently reported to be related with feather pecking. The present study reports the result from a quantitative genetic analysis of feather pecking and three fear test traits in laying hens. Fear was recorded by the tonic immobility test, the open field activity and the emergence box test. These were recorded at a juvenile and adult age of the hens. The heritability of feather pecking was 0.16, and in the range between 0.07 and 0.14 for the fear test traits. Genetic correlations between fear measured in the juvenile and in the adult age point to different but correlated traits. Tonic immobility measured early in life was moderately correlated with feather pecking and might be used as a breeding criterion to reduce feather pecking.

BACKGROUND:Feather pecking (FP) in laying hens is a well-known and multi-factorial behaviour with a genetic background. In a selection experiment, two lines were developed for 11 generations for high (HFP) and low (LFP) feather pecking, respectively. Starting with the second generation of selection, there was a constant difference in mean number of FP bouts between both lines. We used the data from this experiment to perform a quantitative genetic analysis and to map selection signatures.METHODS:Pedigree and phenotypic data were available for the last six generations of both lines. Univariate quantitative genetic analyses were conducted using mixed linear and generalized mixed linear models assuming a Poisson distribution. Selection signatures were mapped using 33,228 single nucleotide polymorphisms (SNPs) genotyped on 41 HFP and 34 LFP individuals of generation 11. For each SNP, we estimated Wright's fixation index (FST). We tested the null hypothesis that FST is driven purely by genetic drift against the alternative hypothesis that it is driven by genetic drift and selection.RESULTS:The mixed linear model failed to analyze the LFP data because of the large number of 0s in the observation vector. The Poisson model fitted the data well and revealed a small but continuous genetic trend in both lines. Most of the 17 genome-wide significant SNPs were located on chromosomes 3 and 4. Thirteen clusters with at least two significant SNPs within an interval of 3 Mb maximum were identified. Two clusters were mapped on chromosomes 3, 4, 8 and 19. Of the 17 genome-wide significant SNPs, 12 were located within the identified clusters. This indicates a non-random distribution of significant SNPs and points to the presence of selection sweeps.CONCLUSIONS:Data on FP should be analysed using generalised linear mixed models assuming a Poisson distribution, especially if the number of FP bouts is small and the distribution is heavily peaked at 0. The FST-based approach was suitable to map selection signatures that need to be confirmed by linkage or association mapping.