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Both humans and nonhuman animals need to show self-control and wait for a larger or better reward instead of a smaller or less preferred but instant reward on an everyday basis. We investigated whether this ability undergoes ontogenetic development in domestic pigs (similar to what is known in human infants) by testing if and for how long nine- and 16-week-old pigs wait for a larger amount of their preferred reward. In a delay-of-gratification task, animals first learned that a small reward was hidden under a white cup and a large reward under a black cup, and then the delay to deliver the large reward was gradually increased. The results show that older pigs could wait longer for a larger reward than younger pigs (10.6 ± 1.3 s vs. 5.2 ± 1.5 s), thereby confirming our hypothesis of ontogenetic development of self-control in pigs. This self-control is likely to be regulated by the behavioural inhibition system and associated systems. Self-control or, more specifically the lack of it may be involved in the development of abnormal behaviours, not only in humans but also in animals. Therefore, research on self-control in decision-making might provide a new perspective on abnormal behaviours in captive animals.

The hypothalamic-pituitary-adrenal (HPA) axis is an important component of neuroendocrine stress regulation and coping behavior. Transcriptome profiles of the hypothalamus and adrenal gland were assessed to identify molecular pathways and candidate genes for coping behavior in pigs. Ten each of high- (HR) and low- (LR) reactive pigs (n = 20) were selected for expression profiling based haplotype information of a prominent QTL-region on SSC12 discovered in our previous genome-wide association study (GWAS) on coping behavior. Comparing the HR and LR pigs showed 692 differentially expressed genes (DEGs) in the adrenal gland and 853 DEGs in the hypothalamus, respectively. Interestingly, 47% (17 out of 36) of DEGs found in both tissues were located in GWAS regions identified on SSC12, indicating that there are significant functional positional candidate genes for coping behaviour. Pathway analysis assigned DEGs to glucocorticoid receptor signaling in the adrenal gland. Furthermore, oxidative phosphorylation, mitochondrial dysfunction, and NGF signaling as well as cholecystokinin/Gastrin-mediated were identified in the hypothalamus. We narrowed the list of candidate genes in GWAS regions by analyzing their DEGs in the HPA axis. The top identified transcripts, including ATP1B2, AURKB, MPDU1 and NDEL1 provide evidence for molecular correlates of coping behavior in GWAS regions.

Emotions, unlike mood, are short-lived reactions associated with specific events. They can be characterized by two main dimensions, their arousal (bodily activation) and valence (negative versus positive). Knowledge of the valence of emotions experienced by domestic and captive animals is crucial for assessing and improving their welfare, as it enables us to minimize the negative emotions that they might experience and to promote positive ones. Emotions can affect vocalizations directly or indirectly through the brain, lungs, larynx or vocal tract. As a result, vocal expression of emotions has been observed across species, and could serve as a non-invasive and potentially very reliable tool to assess animal emotions. In pigs (Sus scrofa), vocal expression of emotions has been relatively well studied. However, it is not known if the vocal indicators revealed in previous studies are valid across call types and contexts. To find this out, we conducted a meta-analysis of the effects of emotional valence on pig vocalizations, including calls recorded in the most common emotional situations encountered by pigs throughout their lives, from birth to slaughter. Our analyses revealed that pigs produced calls characterized by a higher center of gravity, a shorter duration, less noise (lower Wiener entropy), lower formants (measured using the formant dispersion) and LPC coefficients in positive compared to negative contexts. Overall, these vocal parameters could be very useful for developing automated methods to monitor pig welfare on-farm.

The first aim of this study was to establish a surgical procedure to implant a new telemetric device for the continuous recording of electrocardiogram (ECG) and blood pressure (BP) in freely moving pigs. A second aim was the functional assessment of cardiovascular parameters, including heart rate variability (HRV) and blood pressure variability (BPV), so that these data could be used as the basis for the objective evaluation of autonomic activity and balance in different behavioural contexts. Eleven domestic pigs (German Landrace) underwent surgery for the placement of a telemetric device. At day 15 after surgery, 512 consecutive inter-beat intervals and pressure waves were analysed using different detection methods (automatic and manually corrected) while the animals were resting or feeding, respectively. HRV and BPV were calculated. Incomplete datasets were found in four pigs due to missing ECG or BP signals. Technical and surgical issues concerning catheterisation and detachment of the negative ECG lead were continuously improved. In the remaining pigs, excellent signal quality (manually corrected data of 1%) was obtained during resting and acceptable signal quality (

Stress response and coping behavior in pigs are largely shaped by hypothalamic-pituitary-adrenal axis and sympatho-adrenomedullary system action. However, the dynamic interaction between amygdala and hippocampus crucially modulates the behavioral response towards significant emotional events. While this functional relationship is well documented, the molecular underpinnings still remain insufficiently understood. Our study used transcriptome profiling of porcine amygdala and hippocampus to identify molecular pathways that are differentially activated depending on the haplotype of a significantly coping behavior-associated region on pig chromosome 12 (SSC12). The pigs were classified into two groups based on the haplotype information of this QTL-region discovered in our previous genome-wide association study. Ten each of high- (HR) and low- (LR) reactive pigs (n = 20) were selected for differential gene expression analysis and weighted gene co-expression analysis with subsequent pathway analysis. Differentially expressed genes identified in the amygdala include SELL, CXCR7 and NTS, while TRAF3, PTGS2 and CFI were detected in the hippocampus indicating a role of neuroinflammation and immunological processes. Pathway analysis revealed IL-8 signaling, NF-κB signaling, glutamate and GABA metabolism, glucocorticoid receptor signaling and chemokine signaling in the amygdala and ephrin receptor signaling, as well as NF-κB signaling in the hippocampus. We discovered candidate genes in regions detected by genome-wide association study including ARRB2, ADRBK2, THRB, NEK7 and ACVR2B, which relate to dopaminergic and other monoaminergic neurotransmitter systems, neuroimmunomodulation, neuroinflammation and GABA-ergic neurotransmission. These findings provide insights into the molecular underpinning of divergent coping behavior and associated haplotypes in limbic forebrain system in pig.

Cognitive enrichment, a special form of environmental enrichment, addresses the cognitive abilities of animals in captivity. Through cognitive interaction with the environment, the animals regain a certain control over their environment, and essential resources, such as food or water, act as a reward for successful coping. It is assumed that this process has important implications for animal welfare, especially in the intensive housing systems of farm animals. This study investigates the effects of cognitive enrichment on welfare-relevant behaviour (agonistic interactions and behavioural reactivity in a repeated open-field test) and autonomic control (heart rate variability during feeding, resting and in a repeated open-field test) in domestic pigs. A total of forty-eight pigs, Sus scrofa, were housed in groups of four. In six replicates, an experimental group was compared with a conventionally fed control group. The pigs in the experimental group were confronted with a cognitive challenge that was integrated into their familiar housing environment. Pigs were rewarded with food after they successfully mastered the discrimination of an individual acoustical signal followed by an operant task. The pigs in both groups reacted with sympathetic arousal to feeding announcement (increased heart rate (HR)). During feeding, the experimental pigs' HR decreased, and heart rate variability (HRV) increased, while the control pigs' HR stayed highly elevated and HRV decreased. These results are supported by a considerably larger number of agonistic interactions during feeding in the control group. During resting, the basal HRV of the experimental pigs increased (during operant conditioning) compared to the control. In the repeated open-field test, the experimental pigs displayed less locomotion and elimination as well as more contact with the wall and an unknown object compared to the control group. We conclude that cognitive enrichment leads to relaxed feeding and evokes longer lasting positive emotions. Moreover, the pigs displayed more explorative and less fearful behaviour in stressful situations. These findings support the use of cognitive enrichment to improve animal welfare.

Exposure to psychosocial stress can have a profound impact on immune reactivity and health mediated by hypothalamic–pituitaryadrenal (HPA) axis activation. However, current knowledge regarding the mechanisms involved in cross-sensitization between stress and the immune system is limited. Here, we investigated the effects of a single social isolation followed by repeated oral Escherichia coli (E. coli) applications on cortisol, tumor necrosis factor-α (TNF-α), interleukin-6 (IL-6), haptoglobin and C-reactive protein (CRP) concentrations in the blood; on clinical signs of disease; and on mRNA expression of the glucocorticoid receptor (GR), mineralocorticoid receptor (MR), 11β-hydroxysteroid dehydrogenase 1 and 2 (11β-HSD1 and 11β-HSD2), TNF-α and IL-6 in the hypothalamus, prefrontal cortex (PFC) and spleen of 7-, 21- and 35-day-old piglets. Additionally, the protein levels of splenic TNF-α and IL-6 were analyzed. Non-isolated, E. coli-challenged piglets served as a control. Social isolation for 4 h induced a rise in the plasma cortisol concentrations immediately after social treatment and after repeated E. coli applications in isolated compared to non-isolated piglets. The circulating TNF-α concentration was not affected by social treatment. Furthermore, previously isolated piglets showed a higher frequency of signs of disease in response to E. coli challenge than non-isolated piglets, while the haptoglobin and CRP concentrations did not significantly differ between social treatments. In the brain, 11β-HSD1, 11β-HSD2 and IL-6 mRNA expression in the hypothalamus and GR, and 11β-HSD1 and 11β-HSD2 mRNA expression in the PFC were higher in isolated, E. coli-challenged piglets than in the corresponding controls. Moreover, isolated piglets also displayed higher MR, 11β-HSD1 and IL-6 mRNA expression levels and TNF-α concentrations in the spleen. The stress-induced alterations in the hypothalamus and spleen were particularly pronounced in younger piglets. The present findings may contribute to a better understanding of the complex interplay between early psychological stress and an increased risk of disease and might also have implications on aspects of the health and welfare of farm animals and humans.

There is growing evidence that social support given by a conspecific attenuates stress responses of a socially deprived animal. We hypothesized that the presence of a familiar social partner modulates the effectiveness of social buffering as assessed by an altered glucocorticoid sensitivity of immune cells. The current study investigated the effects of a 4-h social deprivation procedure on stress hormone responses and immune cell functions in 7-, 21- and 35-day-old piglets (52 males and 56 females). Within each of the three age categories, nine piglets were deprived of their mother and littermates either alone or with a familiar or unfamiliar age-matched piglet. Compared to non-deprived controls, piglets that were alone displayed significant increases in plasma adrenocorticotropic hormone and cortisol concentrations, and all socially deprived piglets showed a greater in vitro proliferative response of peripheral blood mononuclear cells (PBMCs) to lipopolysaccharide (LPS)-stimulation than controls. Concanavalin A (ConA)-induced in vitro proliferation was not affected by social treatment. Additionally, both the ConA- and LPS-stimulated PBMCs from piglets that experienced any of the deprivation treatments were more resistant to the inhibitory effects of cortisol than PBMCs from the controls in each of the three age categories. Irrespective of the mitogen used, the presence of an age-matched conspecific during deprivation attenuated the dose-dependent cortisol resistance. Here, familiarity between the piglets clearly improved the effectiveness of social support in an age-dependent manner. Collectively, our findings emphasize the benefits of social partners regarding stress coping abilities, with positive implications for animal welfare and health.

The aim of the study was to determine the effects of a single social isolation (4h) of piglets on immediate changes in stress hormones and immune responses at 7, 21 or 35 days of age. This social stressor caused an increase in plasma ACTH and cortisol concentrations and a decrease in plasma TNF-alpha. The percentage of CD8+ cells increased and the CD4+ cell percentage decreased, resulting in decreasing CD4+/CD8+ ratio. The observed changes were consistent for all days studied. Further, the isolation treatment resulted in diminished LPS-stimulated IL-1beta and IL-10 production in whole-blood cultures. In isolated piglets, positive correlations were estimated between changes in percentage of CD8+ cells and cortisol, and negative ones between changes in plasma TNF-alpha and culture supernatant IL-1beta with ACTH and cortisol. The data suggest that psychosocial stress in neonatal pigs induced immune alterations to maintain adaptive stability, but reflecting also negative emotions experienced by this treatment.

Stressful early life events can have short- and long-term effects on neuroendocrine and behavioural mechanisms of adaptation. Here, we investigated the effects of a single social isolation (4 h) of domestic piglets on both behavioural alterations in open-field tests and modifications in the expression of genes regulating glucocorticoid response in stress-related brain regions at 7, 21 or 35 days of age. The mRNAs of glucocorticoid receptor (GR), mineralocorticoid receptor (MR), 11ss-hydroxysteroid dehydrogenase 1 and 2 (11ss-HSD1 and 11ss-HSD2) and c-fos were analysed by real-time RT-PCR in the hypothalamus, hippocampus and amygdala. The social isolation caused both elevated stress hormone concentrations (e.g. cortisol) and open-field reactivity (e.g. locomotion, vocalisation) compared to control piglets. The enhanced behavioural and neuroendocrine activity was associated with distinct changes in gene expression in the limbic system. The hypothalamic GR, MR and 11ss-HSD1 mRNA expressions and the hippocampal 11ss-HSD1 mRNA was significantly higher in isolated piglets, whereas in the amygdala social isolation caused a significant decrease in MR mRNA expression. Isolated piglets also displayed significantly higher c-fos mRNA expression, an estimate of neuronal activation, in hypothalamus and amygdala. The mRNA alterations as well as the behavioural and hormonal pattern show an effect of social isolation on days 7 and 21, but no effect on day 35. In conclusion, a single social isolation in piglets caused age-dependent neuroendocrine and behavioural changes that indicate increased arousal and experienced distress. The present results also suggest that psychosocial stress effects should be considered for the assessment of livestock handling practices with respect to health and welfare.

In the study of animal emotions, emotional valence has been found to be difficult to measure. Many studies of farm animals' emotions have therefore focussed on the identification of indicators of strong, mainly negative, emotions. However, subtle variations in emotional valence, such as those caused by rather moderate differences in husbandry conditions, may also affect animals' mood and welfare when such variations occur consistently. In this study, we investigated whether repeated moderate aversive or rewarding events could lead to measurable differences in emotional valence in young, weaned pigs. We conditioned 105 female pigs in a test arena to either a repeated startling procedure (sudden noises or appearances of objects) or a repeated rewarding procedure (applesauce, toy and straw) over 11 sessions. Control pigs were also regularly exposed to the same test arena but without conditioning. Before and after conditioning, we measured heart rate and its variability as well as the behavioural reactions of the subjects in the test arena, with a special focus on detailed acoustic analyses of their vocalisations. The behavioural and heart rate measures were analysed as changes compared to the baseline values before conditioning. A limited number of the putative indicators of emotional valence were affected by the conditioning. We found that the negatively conditioned pigs showed changes that were significantly different from those in control pigs, namely a decrease in locomotion and an increase in standing. The positively conditioned pigs, however, showed a stronger increase in heart rate and a smaller decrease in SDNN (a heart rate variability parameter indicating changes in autonomic regulation) compared to the controls. Compared to the negatively conditioned pigs, the positively conditioned pigs produced fewer vocalisations overall as well as fewer low-frequency grunts but more high-frequency grunts. The low-frequency grunts of the negatively conditioned pigs also showed lower frequency parameters (bandwidth, maximum frequency, 25% and 50% quartiles) compared to those of the positively conditioned pigs. In any of the statistically significant results, the conditioning accounted for 1.5-11.9% of variability in the outcome variable. Hence, we conclude that repeated moderate aversive and rewarding events have weak but measurable effects on some aspects of behaviour and physiology in young pigs, possibly indicating changes in emotional valence, which could ultimately affect their welfare. The combination of ethophysiological indicators, i.e., the concurrent examination of heart rate measures, behavioural responses and especially vocalisation patterns, as used in the current study, might be a useful way of examining subtle effects on emotional valence in further studies.

Animal personality and coping styles are basic concepts for evaluating animal welfare. Struggling response of piglets in so-called backtests early in life reflects their coping strategy. Behavioral reactions of piglets in backtests have a moderate heritability, but their genetic basis largely remains unknown. Here, latency, duration and frequency of struggling attempts during one-minute backtests were repeatedly recorded of piglets at days 5, 12, 19 and 26. A genome-wide association study for backtest traits revealed 465 significant SNPs (FDR ≤ 0.05) mostly located in QTL (quantitative trait locus) regions on chromosome 3, 5, 12 and 16. In order to capture genes in these regions, 37 transcripts with significant SNPs were selected for expressionQTL analysis in the hypothalamus. Eight genes (ASGR1, CPAMD8, CTC1, FBXO39, IL19, LOC100511790, RAD51B, UBOX5) had cis- and five (RANGRF, PER1, PDZRN3, SH2D4B, LONP2) had trans-expressionQTL. In particular, for PER1, with known physiological implications for maintenance of circadian rhythms, a role in coping behavior was evidenced by confirmed association in an independent population. For CTC1 a cis-expression QTL and the consistent relationship of gene polymorphism, mRNA expression level and backtest traits promoted its link to coping style. GWAS and eQTL analyses uncovered positional and functional gene candidates for coping behavior.

Behavioural patterns and plasma adrenaline, noradrenaline and cortisol responses were studied in domestic pigs with different dominance status during 10-h social confrontation tests with a familiar and an unfamiliar group. Eight trials were carried out, where in each trial two groups of nine growing pigs (12 weeks old) were randomly formed. The pigs with the highest (HR) and lowest (LR) rank from each group were selected as test animals, provided with surgically implanted catheters and kept in single housing for 2 to 3 weeks. After this period, each test animal was introduced once into the familiar and once into the unfamiliar group for 10 h. Introduction of the test animals into the groups caused frequent agonistic interactions during the first 30 min and significantly more agonistic interactions during the confrontation with the unfamiliar group. The agonistic behaviour was accompanied by a rapid increase of plasma catecholamines and cortisol. HR pigs showed significantly higher plasma catecholamine concentrations and more agonistic interactions during the first 30 min compared with the LR pigs. During confrontation with the unfamiliar group, HR pigs experienced more defeats and showed a higher increase of plasma cortisol levels than during the confrontation with the familiar group. No influences of rank or familiarity were found on the other behavioural patterns examined. The results show that agonistic behaviour and activation of the sympatho-adrenomedullary and the hypothalamic-pituitary-adrenal system in pigs during a social confrontation test are determined by the former dominance rank of the animals and the familiarity of the group.

Social deprivation is a severe stressor affecting a number of behavioral and physiological functions of gregarious species. It is assumed that, dependent upon the level of familiarity, social support given by a conspecific may attenuate the adverse consequences of stress. We investigated the effects of a 4h maternal and littermate deprivation on behavioral reactions, stress hormone responses and brain corticosteroid receptor expression in 7-, 21- and 35-day-old domestic piglets (Sus scrofa) that were left alone or in the presence either of a familiar or unfamiliar age-matched piglet. Compared to control animals, all of the socially deprived piglets showed significant stress responses, such as impaired habituation in repeated open-field/novel-object tests, enhanced ACTH and cortisol release, and altered corticosteroid receptor expression in the hypothalamus. In addition, our results demonstrated that younger piglets had more difficulty coping with stress. The presence of an age-matched conspecific had a direct calming effect on the deprived piglet during the deprivation procedure, which was revealed by diminished stress-induced HPA activity and altered reactions in the behavioral test situations (e.g., activity, escape, and vocalization). Furthermore, because the presence of a familiar piglet causes a more pronounced buffering effect, we have shown for the first time that the degree of familiarity between the piglets may influence the effectiveness of social support. Our study emphasizes the benefits of social partners on positive welfare and the ability for pigs to cope with stress; therefore, our results should be taken into account during handling practices such as weaning and mixing.

Animals in the wild are facing a wide variety of challenges and ever-changing environmental stimuli. For successful coping, animals use both innate behavioural programs and their cognitive skills. In contrast, zoo- and farm animals have to cope with restricted husbandry conditions, which offer only few opportunities to adequately satisfy their various needs. Consequences could be sensory and cognitive underchallenge that can cause boredom and frustration as well as behavioural disturbances. Initially intended for improvement of management and husbandry, different forms of operant behavioural training have been applied firstly in zoo- and later also in farm animals. It has been suggested that successful coping with appropriate cognitive challenges is a source of positive emotions and may lead to improved welfare. Under the term cognitive enrichment, new approaches have been developed to integrate cognitive challenges into the housing of zoo- and farm animals. The present article reviews actual research in the field. Previous results indicate that, beyond improvement of management and handling routines, such approaches can positively affect animal behaviour and welfare. The combination of explorative and appetitive behaviour with successful learning improves environmental predictability and controllability for the animals, activates reward-related brain systems and can directly affect emotional processes of appraisal. For practical implementation in farm animal husbandry, it sounds promising to link individual access to e.g. automated feeders or milking systems with previously conditioned stimuli and/or discriminatory learning tasks. First experimental approaches in pigs, dwarf goats and cattle are available and will be discussed in the present article.

Weaning of farm animals is of biological and economical relevance. Due to economical considerations, the weaning age of piglets (Sus scrofa) has drastically been reduced during the last few years, whereas biological consequences remain ambiguous. This review gives a survey of current international research on weaning and its psychobiological consequences in conventional pig farming. The influence of weaning age on behaviour, neuroendocrine and immune systems of piglets is the main focus of this paper. The abrupt weaning in pig farming comprises multiple stressful events such as change in diet, new physical environment, as well as the often underestimated psychosocial consequences of maternal deprivation and regrouping with unfamiliar conspecifics. Results presented in this review suggest that early weaned piglets (or = 21 days of age) show enhanced behavioural problems and neuroendocrine stress reactions as well as reduced immune responses. These adaptation problems may have short- and long-term effects on animals' welfare and health, consequently resulting in increased financial requirements and higher management demands. Weaning before the third week of age is not recommended. Additionally, alternative housing systems (e.g. group housing) with adequate weaning methods (e.g. weaning without regrouping, social contacts prior to weaning) to diminish psychosocial stress are discussed.

Psychosocial stress in the form of maternal deprivation and social isolation during early postnatal life induces persistent alterations in behavioral and physiological mechanisms of adaptation. One consequence may be an increased susceptibility to diseases in later life. Therefore, the aim of the present study was to investigate in domestic piglets the effects of a repeated social isolation (2 h daily from day 3 to day 11 of age) on behavioral, endocrine and immune responses to an endotoxin challenge with lipopolysaccharide (LPS) 1 day or 45 days after the isolation period. Peripheral LPS administration caused serious sickness behavior (somnolence, shivering, vomiting) and provoked profound increases in circulating tumor necrosis factor-alpha (TNF-alpha), ACTH and cortisol concentrations. The prior social isolation treatment enhanced signs of sickness and impaired suckling behavior. Early isolated piglets responded to LPS by an increase of shivering on day 12 and by increased vomiting on day 56 compared to controls. Further, there were considerable delays and reductions of time isolated piglets spent suckling on day 12. The repeated isolation stressor diminished TNF-alpha increases after LPS, whereas stress hormone levels were not significantly affected by isolation treatment. Finally, stronger relationships between signs of sickness and physiological measures were revealed in early isolated piglets. The duration of somnolence in isolated piglets was related to changes of cortisol and TNF-alpha concentrations, and the highest impact on duration of shivering was found for changes in cortisol and corticosteroid binding globulin levels. The present results suggest a sustained adaptive sensitization of coping with infection by social stress experience during early development in piglets.

The composition of sow colostrum and milk from single teats were characterized in 20 primiparous German Landrace sows at ten points throughout lactation time (after birth of the first piglet, and subsequently at 12 and 24 h, 3, 5, 7, 14, 21, 28 and 35 d postpartum). In these samples contents of protein, IgG, lactose and fat were analyzed. In the first 3 days of lactation protein- and IgG-contents drastically decreased, on the other hand the levels of energy resources lactose and fat increased. Significant effects of the single teat and the teat pair on the content of protein at farrowing and 12 h later as well as on the contents of IgG and lactose 12 h postpartum were found. The highest contents of protein and IgG were analyzed in colostrum of teats in the median udder section, the colostrum of cranial teats had the highest content of lactose. At days 3 and 5 of lactation milk showed large coefficients of variation (cv) in milk composition from teat to teat as well as from sow to sow (intraindividual cv: protein: 22%, IgG: 45%, lactose: 20%, fat: 34%; interindividual cv: protein: 36%, IgG: 85%, lactose: 27%, fat: 47%). During lactation period the intraindividual variation of milk parameters was lower than the interindividual variation. Possible reasons for these results are discussed.

The effects of an enriched environment on growth, meat quality, and muscle biological traits were investigated using 64 castrated pigs (age range 7 to 20 weeks). Positive emotional appraisal was induced by a system that acoustically frequently called individuals out of a group (n=32) to a feeding station, where they were rewarded with small portions of feed. The results were compared with traditional fed pigs (Control; n = 32). There was no environmental effect on live weight and lean muscle percentage. However, the stimulating keeping regime reduced the intramuscular fat content of the M. longissimus by 0.2 % and increased the protein content by 0.5 %. Moreover, the drip loss of M. longissimus was 1.1 % lower in the treatment group. The analysis of muscle fibre traits (biopsy samples of M. longissimus) showed a 7 % higher proportion of oxidative muscle fibres in the pigs reared in the enriched environment. This fibre type showed a tendency to increased fibre areas. These findings indicate a higher oxidative capacity of the muscle. However, the activity of the inarker enzyme citrate synthase (CS) was not influenced and the activity of lactate dehydrogenase (LDH) was even increased compared to the control animals.

Social stress during early postnatal life often results in long-term effects on neuroendocrine and immune adaptation mechanisms. Therefore, the aim of the present study was to determine the influence of a 2-h daily social isolation from Day 3 to Day 11 on the acute and long-term proinflammatory and neuroendocrine responses of piglets challenged with the bacterial endotoxin lipopolysaccharide (LPS; 100 microg/kg body weight). Peripheral LPS administration significantly increased plasma concentrations of tumor necrosis factor-alpha (TNF-alpha), ACTH and cortisol in isolated and control pigs. However, the activity of the hypothalamic-pituitary-adrenal (HPA) axis after LPS stimulation was not significantly affected by isolation treatment, whereas the prior social isolation diminished the plasma TNF-alpha response to LPS 1 day as well as 45 days after the isolation period. The hippocampal TNF-alpha concentration in response to LPS was also reduced in priorly isolated pigs compared to control animals. Furthermore, the significant increase of TNF-alpha in the spleen caused by LPS was associated with a dramatic decrease in glucocorticoid receptor (GR) binding. The GR binding in hippocampus was increased in isolated pigs and was significantly decreased after LPS injection. In addition, the repeated isolation stressor was shown to increase hippocampal levels of interleukin-1beta (IL-1beta). The present results indicate that repeated social isolation of neonatal pigs may cause long-term effects on proinflammatory regulation at the periphery and in the brain following immune challenge with particular importance of TNF-alpha in mediating these interactions.

The nursing and suckling behaviour of the domestic pig is a complex social behavioural interaction aimed to ensure the milk flow from the sow to the piglets. Ultimately caused theoretical frameworks from basic ethology and/or sociobiology are increasingly used to explain such social behavioural patterns. The present paper gives a short survey of ethophysiological aspects of the nursing and suckling behaviour in domestic pigs and, thereafter, discusses the development and consequences of some parts of the suckling behaviour and the maternal lactation performance in terms of a possible parent-offspring and weaning conflict. It is concluded that--additional to proximately caused mechanisms--the ontogeny of domestic pigs can be interpreted with such principles. In housing systems allowing the animals more control over their behaviour (e.g. their investment) ultimate causes should be more considered. Especially the pathways on which the ultimate strategies become effective via proximate mechanisms do not seem to be understood in total and require specific research. However, it is suggested that effective improvements of welfare, health and productivity can only be reached with the knowledge and integration of both ultimate and proximate causes of behaviour.

The impact of a 10-h social confrontation on behavior, plasma adrenaline, noradrenaline and cortisol was studied in 14 domestic pigs (eight castrated males and six females) with prior dominance experience. Prior to the experiments, animal groups, each consisting of nine growing pigs (12 weeks old) from different litters, were composed randomly. After 5 days, the pig with the highest rank from each group was removed, provided with a jugular vein catheter and kept in single housing for 2-3 weeks. After this period, each test animal was returned into its familiar group for a 10-h social confrontation. The reintroduction of the test animals into the familiar groups caused frequent agonistic interactions during the first 30 min. Seven animals succeeded to win most of their encounters during the confrontation test and were classified as High Success (HS) animals, whereas seven other animals lost most of their encounters and were classified as Low Success (LS) animals. The reintroduction of the test animals into the groups provoked also marked changes in behavioral and physiological measures. The frequent fighting behavior during the first 30 min was accompanied by a rapid increase of plasma catecholamines and a delayed increase of cortisol. Immediately after introduction, LS pigs tended to show higher plasma adrenaline and noradrenaline concentrations than HS pigs. There was also a tendency for a sustained increase of noradrenaline/adrenaline ratios in HS pigs, whereas the ratios of LS pigs remained nearly unchanged. No significant differences between both groups were found in cortisol levels and in the frequency of agonistic interactions. However, LS animals showed less locomotion, more lying and spent less time exploring the pen or other animals. These results show that during a social confrontation the experience of defeats for formerly high-ranking pigs is accompanied by increased submissive or passive behavior and a higher sympathoneural and adrenomedullary reaction, which may indicate more emotional distress and fear compared to successful animals.

It is assumed that calls may give information about the inner (emotional) state of an animal. Hence, in the last years sound analysis has become an increasingly important tool for the interpretation of the behavior, the health condition, and the well-being of animals. A procedure was developed that allows the characterization, classification, and visualization of the cluster structures of stress calls of domestic pigs (Sus scrofa). Based on the acoustic model of the sound production the extraction of features from calls was performed with linear prediction coding (LPC). A vector-based self-organizing neuronal network was trained with the determined LPC coefficients, resulting in a feature map. The cluster structure of the calls was then visualized with a unified matrix and the neurons were labeled for their input origin. The basic applicability of the procedure was tested by using two examples which were of special interest for a possible evaluation of the normal farming practice. The procedure worked well both in discriminating individual piglets by their scream characteristics and in classifying pig stress calls vs other calls and noise occurring under normal farming conditions.

Despite technological changes and improved management, piglet mortality remains a problem for both production and welfare. Most preweaning mortality occurs within the first 3 days after birth because of problems with adaptation and development. Thus, the purpose of our study was to determine the physiologic state of newborn pigs with respect to piglet survival. Data were collected from 1024 live-born piglets of 106 primiparous German Landrace sows to analyze relationships between farrowing traits, early postnatal vitality and blood chemistry, including immunity of piglets at birth. Surviving piglets were compared with those that died during the first 10 days of life. The survivors were significantly heavier at birth (P=0.001), were born earlier in the birth order (P=0.04), reached the udder and took in first colostral milk more quickly (P=0.001) and had a smaller drop in rectal temperature I h after birth (P=0.001) than dead. However, dead piglets had significantly higher blood levels of inorganic phosphorus (P=0.0001), calcium (P=0.04) and urea (P=0.05), but a lower concentration of alpha2-macroglobulin and lower lymphocyte proliferation indices in response to pokeweed mitogen (P=0.05). Models fitted for discrimination between survivors and piglets that died included, in addition to birth weight and litter size, the foraging behavior of neonates (time from birth to first suckle) and their thermoregulatory capacity (rectal temperature 1 h after birth) in the first experimental unit, as well as prenursing biochemical measures (inorganic phosphorus, calcium and glucose) in the second experimental unit. These ethophysiological and biochemical traits of early postnatal vitality are important determinants of maturity and development at birth. Hence, breeding programs and perinatal housing and feeding conditions should ensure a high physiological maturity to improve mortality rates of neonates.

The paper presents a new system for the automatic monitoring of open field activity and choice behaviour of medium-sized animals. Passive infrared motion detectors (PID) were linked on-line via a digital I/O interface to a personal computer provided with self-developed analysis software based on LabVIEW (PID technique). The set up was used for testing 18 one-week-old piglets ( Sus scrofa) for their approach to their mother's nursing vocalization replayed through loudspeakers. The results were validated by comparison with a conventional Observer technique, a computer-aided direct observation. In most of the cases, no differences were seen between the Observer and PID technique regarding the percentage of stay in previously defined open field segments, the locomotor open field activity, and the choice behaviour. The results revealed that piglets are clearly attracted by their mother's nursing vocalization. The monitoring system presented in this study is thus suitable for detailed behavioural investigations of individual acoustic recognition. In general, the PID technique is a useful tool for research into the behaviour of individual animals in a restricted open field which does not rely on subjective analysis by a human observer.

The effects of social rank on immune, metabolic, and endocrine responses were studied in 10 newly mixed groups of German Landrace pigs (9 individuals each) at an age of 12 weeks. Immediately after mixing, the agonistic interactions (AI) of all group members were continuously recorded over 3 days (10 h daily). An individual dominance value (DV) was calculated by the number of wins minus defeats in relation to all decisive fights (DVor = 0, subordinate; DV0, dominant). Blood samples were taken 24 h before and 3 days after mixing. The data showed that the social status had a significant effect on lymphocyte proliferation in responses to different mitogens: socially dominant pigs had higher proliferative response than subordinate pigs. In addition, during the observation period the lymphocyte activation by mitogens increased in the dominant animals and decreased in the subordinate animals with increasing number of agonistic interactions. The rise in total serum IgG concentration 3 days after mixing was higher in dominant pigs compared with subordinates. The dominance status did not significantly affect plasma metabolic levels nor cortisol concentrations. However, mixing appeared to increase glucose and total protein values and to decrease alkaline phosphatase and cortisol levels in both, dominant and subordinate pigs. In conclusion, mitogen induced cell proliferation seems to be a valuable marker for acute social stress in pigs.

Investigations with 693 piglets have shown, that suckling piglets with preference of cranial teat pairs were significantly superior to their littermates in all parameters of early postnatal vitality: they stood up and took in first colostrum earlier and showed a lower postnatal drop in rectal temperature in comparison with their siblings. Piglets in cranial teat position had the highest daily gain (205 g, p0.05) during suckling period. Highly significant influences of vitality of piglets post natum on live weight development and mortality up to 28th day of age were shown. The faster piglets took in colostrum after birth and the lower postnatal drop in rectal temperature was, the higher was the obtained daily gain during suckling period. Piglets which died within the first 7 or 28 days of age had needed a significantly longer time till first postnatal colostrum intake and a more pronounced drop in rectal temperature in comparison with their littermates.

The suckling behaviour, the development of daily weight gains and biochemical blood parameters (immunoglobulin G, total protein, glucose, urea) were investigated during the suckling period (35 days) of piglets (Sus scrofa) in a group housing system for nursing sows (group P: 4 primiparous sows, 34 piglets; group M: 4 multiparous sows, 43 piglets). Following an imprinting period of 10 days, piglets were given the opportunity to leave the farrowing pen and to interact with the other sows (variant e: suckling bouts at own mother only, variant f: suckling bouts also at one or more unfamiliar sows). Although a number of piglets suckled at unfamiliar sows (Pf = 73.5%; Mf = 32.5%; p = 0.01), only those piglets that stayed with their own mother after mixing had some "physiological advantages" (Pe: daily weight gain from day 10 to 35, p = 0.05; Me: IgG, p = 0.05). Piglets of primiparous sows had a lower teat pair fidelity within the imprinting period (day 1 to 10). During this time, piglets of both groups suckling at unfamiliar sows showed a lower teat pair fidelity (P: ns; M: p = 0.05), tended to prefer the posterior part of the udder, and their blood glucose and urea contents were higher (P: ns; M: p = 0.05). This was interpreted as an indication of competitive strain which obviously encouraged the piglets to leave their mothers. It is necessary to carry out further systematical studies on sociophysiological mechanisms.

Viewing dominance as an attribute of repeated agonistic interactions between two individuals, the present paper reviews theoretical approaches towards concepts of dominance, methods of measurement, and basic principles and problems connected with social dominance in domestic pigs. Domestic pigs are able to establish social organization structures during all stages of their ontogeny. According to definition, dominance relationships occur when a consistent asymmetry of the result of dyadic agonistic interactions can be assessed. This must not necessarily be connected immediately with a better availability of resources, or a high stability of existing dominance relationships, or a functional definition of dominance. When sociometric characteristics are calculated, it seems to be appropriate to use them for different levels of a biological system (individual, individual pair, group). Investigations of social behaviour and dominance in farm animals should take into account that mechanisms of social behaviour in confined environments are often carried out in parts only. Connections of the dominance concept with other concepts of behavioural regulation should be theoretically considered and further investigated by experimental studies.

The shot biopsy is a common experimental technique for the collection of samples to investigate muscle tissue characteristics or to determine meat quality features in pigs. Its application seems to be also possible in interdisciplinary research projects investigating animal stress, behaviour, and welfare. The present study on 12 group-housed pigs (age: 12 weeks, weight: 29.3 kg) shows the influence of this wound-causing technique on different humoral and cell-mediated parameters of the immune system at 1, 3, 5, and 9 days after biopsy compared to the initially investigated levels before. An enhancement of the blood sedimentation rate and both the IgG and the cellular immune response in vivo (leukocytes, lymphocytes) as well as in vitro (ConA) was observed. Furthermore, there were signs of a secondary wound infection 5 days after biopsies were taken possibly caused by mutual oral manipulation of the animals. We conclude that the muscle shot biopsy technique can be used on group-housed pigs as a method to investigate muscle physiological characteristics. The technique, however, induces immunological reactions which may interfere with stress-induced immune reactions.

Investigations with 1248 newborn piglets in 7 farms showed a high significant influence of birth weight on parameters of early postnatal vitality. The duration between birth and first standing up was by two times, the time between birth and first udder contact by 3.5 times and the duration between birth and first colostrum intake was by 4 times longer in piglets with a low birth weight (800 g) in comparison with heavier piglets at birth (2200 g). The drop in rectal temperature up to 30 minutes after birth reached 4.5 Kelvin in lightweight piglets, whereas their litter mates with a high body weight at birth had a value of 0.85 K (p0.01). The birth weight of piglets belongs to the most important endogenous factors with influence on the vitality of newborn piglets and has a high prognostic value in relation to the risk of losses and the live weight development of neonates.