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It is crucial to identify whether relations between immune characteristics and damaging behaviors in production animals exist, as these behaviors reduce animal welfare and productivity. Feather pecking (FP) is a damaging behavior in chickens, which involves hens pecking and pulling at feathers of conspecifics. To further identify relationships between the immune system and FP we characterized high FP (HFP) and low FP (LFP) selection lines with regard to nitric oxide (NO) production by monocytes, specific antibody (SpAb) titers, natural (auto)antibody (N(A)Ab) titers and immune cell subsets. NO production by monocytes was measured as indicator for innate pro-inflammatory immune functioning, SpAb titers were measured as part of the adaptive immune system and N(A)Ab titers were measured as they play an essential role in both innate and adaptive immunity. Immune cell subsets were measured to identify whether differences in immune characteristics were reflected by differences in the relative abundance of immune cell subsets. Divergent selection on FP affected NO production by monocytes, SpAb and N(A)Ab titers, but did not affect immune cell subsets. The HFP line showed higher NO production by monocytes and higher IgG N(A)Ab titers compared to the LFP line. Furthermore the HFP line tended to have lower IgM NAAb titers, but higher IgM and IgG SpAb titers compared to the LFP line. Thus, divergent selection on FP affects the innate and adaptive immune system, where the HFP line seems to have a more responsive immune system compared to the LFP line. Although causation cannot be established in the present study, it is clear that relationships between the immune system and FP exist. Therefore, it is important to take these relationships into account when selecting on behavioral or immunological traits., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Feather pecking (FP), a serious welfare and economic issue in the egg production industry, has been related to coping style. Proactive and reactive coping styles differ in, among others, the stress response, serotonergic activity and immune activity. Yet, it is unknown whether genetic lines divergently selected on FP (i.e. FP genotypes) or individuals differing in FP (i.e. FP phenotypes) can be categorized into coping styles. Therefore, we determined peripheral serotonin (5-HT) levels, natural antibody (NAb) titers, behavioral and corticosterone (CORT) responses to manual restraint (MR) in FP genotypes (high FP (HFP), low FP (LFP) and unselected control (CON) line) and FP phenotypes (feather pecker, feather pecker-victim, victim and neutral). We further examined the consistency of and relationships between behavioral and physiological measures. FP genotypes differed in behavioral responses to MR, 5-HT levels and NAb titers, but not in CORT levels after MR. HFP birds had less active responses at adolescent age, but more active responses at adult age compared to LFP and CON birds. The CON line had higher 5-HT levels at adolescent age, while the HFP line had lower 5-HT levels than the other lines at adult age. Overall, the HFP line had lower IgM NAb titers, while the LFP line had lower IgG NAb titers compared to the other lines. FP phenotypes differed in behavioral responses to MR and 5-HT levels, but not in CORT levels after MR or NAb titers. Within the HFP line, feather peckers tended to have less active responses compared to neutrals at adolescent age, while victims had more active responses compared to the other phenotypes at adult age. Feather peckers had higher 5-HT levels than neutrals at adult age. Behavioral and CORT responses to MR were not consistent over time, suggesting that responses to MR might not reflect coping style in this study. Furthermore, proactive behavioral responses were correlated with reactive physiological measures and vice versa. Thus, it was not possible to categorize FP genotypes or FP phenotypes into specific coping styles., (Copyright © 2018 Elsevier Inc. All rights reserved.)

The presence of a conspecific during a stressful situation, i.e. social support, can considerably lower the stress response of an individual compared to experiencing the stressful situation alone. Pigs also benefit from social support, but it is not known whether the extent to which they benefit is dependent on their personality or coping style. In this study, therefore, the effect of social support on behavioral and physiological (i.e. salivary cortisol, heart rate and the heart rate variability parameters standard deviation (SDNN) and root mean square of successive differences (RMSSD)) stress responses was studied in pigs with different coping styles. Based on the backtest, piglets were classified as high-resisting (HR) or low-resisting (LR). At 11weeks of age, 12 gilts of each coping style were subjected to a 15min restraint test in a weighing cage without a pen mate present (i.e. no support treatment) and 12 other gilts of each coping style were subjected to this test with a pen mate, a boar with the same coping style, present (i.e. support treatment). With the pen mate present, LR gilts showed less standing alert behavior and they had their ears back less often than without the pen mate present. On the other hand, HR gilts seemed to spend less time on escaping the cage and more HR gilts seemed to urinate in the situation when the pen mate was present than without the pen mate present, but this was not significant. Independently of the test situation, HR gilts grunted more than LR gilts and they were more likely to urinate than LR gilts. Salivary cortisol concentrations were not affected by treatment or coping style, but were increased at 30min after the start of the test after which concentrations decreased again to starting levels. The heart rate and heart rate variability parameters RMSSD and RMSSD/SDNN ratio were not affected by treatment or coping style, but the SDNN was lower in the LR pigs during the first and last 5min of the test when a pen mate was present than without the pen mate present. In addition, heart rate and heart rate variability were increased during the test compared to before and after the test. No differences in behavior and salivary cortisol concentrations were found between the HR and LR accompanying boars. Although effects of social support were not found on all variables measured, the results do indicate that pigs may benefit from social support during a stressful situation and that pigs with a low-resisting coping style will likely benefit more than pigs with a high-resisting coping style., (Copyright © 2014 Elsevier Inc. All rights reserved.)

Severe feather pecking (SFP) in laying hens is a detrimental behavior causing loss of feathers, skin damage and cannibalism. Previously, we have associated changes in frontal brain serotonin (5-HT) turnover and dopamine (DA) turnover with alterations in feather pecking behavior in young pullets (28-60 days). Here, brain monoamine levels were measured in adult laying hens; focusing on four brain areas that are involved in emotional behavior or are part of the basal ganglia-thalamopallial circuit, which is involved in obsessive compulsive disorders. Three behavioral phenotypes were studied: Severe Feather Peckers (SFPs), Victims of SFP, and Non-Peckers (NPs). Hens (33 weeks old) were sacrificed after a 5-min manual restraint test. SFPs had higher 5-HIAA levels and a higher serotonin turnover (5-HIAA/5-HT) in the dorsal thalamus than NPs, with intermediate levels in victims. NPs had higher 5-HT levels in the medial striatum than victims, with levels of SFPs in between. 5-HT turnover levels did not differ between phenotypes in medial striatum, arcopallium and hippocampus. DA turnover levels were not affected by feather pecking phenotype. These findings indicate that serotonergic neurotransmission in the dorsal thalamus and striatum of adult laying hens depends on differences in behavioral feather pecking phenotype, with, compared to non-pecking hens, changes in both SFP and their victims. Further identification of different SFP phenotypes is needed to elucidate the role of brain monoamines in SFP., (© 2013.)

Pigs differ in their behavioural responses towards environmental challenges. Individual variation in maladaptive responses such as tail biting, may partly originate from underlying biological characteristics related to (emotional) reactivity to challenges and serotonergic system functioning. Assessing relations between behavioural responses and brain and blood serotonin parameters may help in understanding susceptibility to the development of maladaptive responses. The objective of the current study was, therefore, to assess the relationship between the pigs' serotonergic parameters measured in both blood and brain, and the behaviour of pigs during a novelty test. Pigs (n=31) were subjected to a novelty test at 11weeks of age, consisting of 5-min novel environment exposure after which a novel object (a bucket) was introduced for 5min. Whole blood serotonin, platelet serotonin level, and platelet serotonin uptake were determined at 13weeks of age. Levels of serotonin, its metabolite and serotonin turnover were determined at 19weeks of age in the frontal cortex, hypothalamus and hippocampus. The behaviour of the pigs was different during exposure to a novel object compared to the novel environment only, with more fear-related behaviours exhibited during novel object exposure. Platelet serotonin level and brain serotonergic parameters in the hippocampus were interrelated. Notably, the time spent exploring the test arena was significantly correlated with both platelet serotonin level and right hippocampal serotonin activity (turnover and concentration). In conclusion, the existence of an underlying biological trait - possibly fearfulness - may be involved in the pig's behavioural responses toward environmental challenges, and this is also reflected in serotonergic parameters., (Copyright © 2013 Elsevier Inc. All rights reserved.)

For the welfare of group-housed animals, such as pigs, the emotional state of an individual pig is relevant, but also the extent to which pen mates are affected by the distress or pleasure of other individuals, i.e. emotional contagion, a simple form of empathy. Therefore, indicators of positive and negative emotions were investigated in pigs during anticipation and experience of a rewarding (access in pairs to a compartment with straw, peat and chocolate raisins) or aversive (social isolation combined with negative, unpredictable interventions) event. Thereafter the same indicators were investigated in naive pigs during anticipation and experience of a rewarding or aversive event by their trained pen mates. Positive emotions could be indicated by play, barks and tail movements, while negative emotions could be indicated by freezing, defecating, urinating, escape attempts, high-pitched vocalizations (screams, squeals or grunt-squeals), tail low, ears back and ear movements. Salivary cortisol measurements supported these behavioral observations. During anticipation of the aversive event, naive pigs tended to show more tail low. During the aversive event, naive pigs tended to defecate more, while they played more during the rewarding event. These results suggest that pigs might be sensitive to emotional contagion, which could have implications for the welfare of group-housed pigs. Pig emotions and the process of emotional contagion merit, therefore, further research., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Fearfulness of an individual can affect its sensitivity to stress, while at the same time the social situation in which an animal lives can affect its fear level. It is however unknown what the long-term effects of high fearfulness on sensitivity to stress are, on individual or group level in laying hens. We hypothesize that increased fearfulness at a young age results in increased sensitivity to stress at an adult age, and that this relation can differ between groups, due to differences in group composition. Therefore, we studied the relation between fearfulness in an Open Field (OF) test at six weeks of age and plasma-corticosterone (CORT) levels after a 5-min Manual Restraint test (MR) at 33 weeks of age, and assessed behavior in the home pen. We used birds from a low mortality line, selected for four generations on low mortality due to feather pecking and cannibalism and a control line (n=153 in total, eight pens/line). These lines are known to differ in fearfulness and stress physiology. Chicks from the low mortality line were more active in the OF compared to chicks from the control line. Chicks that showed a fearful response (no walking, no vocalizing) in the OF test had higher CORT at 33 weeks of age than chicks that walked and/or vocalized in the OF test and had higher activity in the home pen as adults. On group level, a passive response in the OF was related to high CORT levels after MR. Presence of at least one fearful bird in a group led to higher CORT in the other group mates compared to birds from groups with no fearful birds present. Birds from groups in which more than 50% of birds had severe comb lesions had higher CORT levels compared to birds from groups with less than 50% of birds affected. High fearfulness of laying hen chicks can on individual level have a long-term effect on stress sensitivity. The presence of fearful birds in a group as well as signs of social instability in a group, indicated by comb lesions, can affect sensitivity to stress of birds from the same group. The mechanism by which this occurs can lie in social transmission of (fear related) behavior, but this suggestion needs further investigation., (Copyright © 2012 Elsevier Inc. All rights reserved.)

The aim of the present study was to investigate the effect of brooding and group selection for low mortality on post-stress corticosterone and peripheral serotonin in laying hens. Birds in the experiment originated from the same population and were either group-selected for low mortality (low mortality line) or randomly selected (control line) for two generations. Twelve groups of seven birds from each line were used. Within each line, six groups were brooded by a foster mother and six groups were non-brooded. At 33 weeks of age, birds (n=42/treatment) were manually restrained for 5 min, during which their behavioral response (number of struggles) was studied. Fifteen minutes after the start of the manual restraint, blood samples were drawn for assessment of plasma corticosterone and whole blood serotonin (5-HT) concentration. In the low mortality line, 80% of the birds struggled and vocalized vs. 72% in the control line (non significant). Birds from the control line had a higher plasma corticosterone concentration after manual restraint than birds from the low mortality line (7.7 vs. 6.0 nmol ml(-1)). Furthermore, birds from the control line that were reared without a mother had a lower whole-blood 5-HT concentration than birds from the other treatments (45 vs. 48 nmol ml(-1)). These results indicate that both brooding and selection for low mortality affect post-stress corticosterone and peripheral serotonin concentration, which may result in a reduced propensity to develop feather pecking.

Severe feather pecking is a maladaptive behavior in laying hens that may result in cannibalism and ultimately death of the victims. Selection methods in which the genetic effect of an animal on the survival of its group members is taken into account, i.e. 'group selection', have been shown to be very effective in reducing mortality due to feather pecking and cannibalism in laying hens. It has been suggested that fearfulness and serotonergic functioning are involved in the causation and development of feather pecking. We investigated effects of selection based on survivability in non-beak trimmed groups on fear-related behavior and peripheral 5-HT concentration and uptake in hens with trimmed or intact beaks, in a 2 x 2 factorial arrangement. Hens from the second generation of the low mortality line showed less fear-related behavior than control line hens. In addition, they displayed higher whole blood 5-HT concentrations and a lower platelet 5-HT uptake, indicating differences in functional activity of the 5-HT system. Beak trimming resulted in lowered levels of fear, and in a reduction of feather damage. Hens with trimmed and intact beaks did, however, not differ in peripheral 5-HT. The results imply that fearfulness and 5-HT activity are related to feather pecking without distinguishing between cause and effect. However, given that selection altered fear-related behaviors and peripheral 5-HT concentration and uptake, whereas beak trimming affected fearfulness and feather damage, but not 5-HT parameters, we suggest that peripheral 5-HT activity might reflect the predisposition to develop severe feather pecking under adverse conditions in laying hens.