H.J. Blokhuis

Coping styles in animals: current status in behavior and stress-physiology

This paper summarizes the current views on coping styles as a useful concept in understanding individual adaptive capacity and vulnerability to stress-related disease. Studies in feral populations indicate the existence of a proactive and a reactive coping style. These coping styles seem to play a role in the population ecology of the species. Despite domestication, genetic selection and inbreeding, the same coping styles can, to some extent, also be observed in laboratory and farm animals. Coping styles are characterized by consistent behavioral and neuroendocrine characteristics, some of which seem to be causally linked to each other. Evidence is accumulating that the two coping styles might explain a differential vulnerability to stress mediated disease due to the differential adaptive value of the two coping styles and the accompanying neuroendocrine differentiation.

Housing familiar male wildtype rats together reduces the long-term adverse behavioural and physiological effects of social defeat.

Social stress in rats is known to induce long-lasting, adverse changes in behaviour and physiology, which seem to resemble certain human psychopathologies, such as depression and anxiety. The present experiment was designed to assess the influence of individual or group housing on the vulnerability of male Wildtype rats to long-term effects of inescapable social defeat. Group-housed rats were individually exposed to an aggressive, unfamiliar male conspecific, resulting in a social defeat. Defeated rats were then either individually housed or returned to their group. The changes in their behaviour and physiology were then studied for 3 weeks. Results showed that individually housed rats developed long-lasting, adverse behavioural and physiological changes after social defeat. Their body growth was significantly retarded (p < .05) between 7 and 14 days after defeat. When individually and group-housed rats were exposed to a mild stressor (sudden silence) 2 days after defeat, both groups became highly immobile. However, when exposure was repeated at day 21, individually housed rats were still highly immobile compared to group-housed rats which regained their normal mobility after only 7 days. In an open field test, also regularly repeated, individually housed rats took significantly longer to leave their home base and were also significantly less mobile than group-housed rats over the entire 3-week test period as well as at specific timepoints. When the rats were placed in an elevated plus-maze 14 days after defeat, those that were individually housed were significantly more anxious than those that were group-housed. When tested at 21 days after defeat in a combined dexamethasone (DEX)/corticotrophin-releasing factor (CRF) test, results showed that the hypothalamic-pituitary-adrenocortical (HPA) activity in individually housed rats was higher. This was evidenced in the latter animals by the fact that DEX was significantly less able to suppress the secretion of ACTH and corticosterone, and by a significantly higher release of ACTH after administration of CRF. Although the weights of the spleen and testes of the two groups did not differ, the adrenals of individually housed rats were larger and the thymus and seminal vesicles were smaller. We conclude that when rats are isolated after defeat, they show long-lasting, adverse behavioural and physiological changes that resemble symptoms of stress-related disorders. In contrast, when familiar rats are housed together these effects of a social defeat are greatly reduced. These findings show that housing conditions importantly influence the probability of long-term adverse behavioural and physiological effects of social defeat in male Wildtype rats.

Heart rate and heart rate variability during a novel object test and a handling test in young horses.

Forty-one Dutch Warmblood immature horses were used in a study to quantify temperamental traits on the basis of heart rate (HR) and heart rate variability (HRV) measures. Half of the horses received additional training from the age of 5 months onwards; the other half did not. Horses were tested at 9, 10, 21 and 22 months of age in a novel object and a handling test. During the tests, mean HR and two heart variability indices, e.g. standard deviation of beat-to-beat intervals (SDRR) and root mean square of successive beat-to-beat differences (rMSSD), were calculated and expressed as response values to baseline measures. In both tests, horses showed at all ages a significant increase in mean HR and decrease in HRV measures, which suggests a marked shift of the balance of the autonomic nervous system towards a sympathetic dominance. In the novel object test, this shift was more pronounced in horses that had not been trained. Furthermore, statistical analysis showed that the increase in mean HR could not be entirely explained by the physical activity. The additional increase in HR, the nonmotor HR, was more pronounced in the untrained horses compared to the trained. Hence, it is suggested that this nonmotor HR might be due to the level of emotionality. HR variables showed consistency between years, as well as within the second year. These tests bring about a HR response in horses, part of which may indicate a higher level of emotionality; and horses show individual consistency of these HR variables over ages. Therefore, it is concluded that mean HR and HRV measures used with these tests quantify certain aspects of a horses temperament.

Feather-pecking in poultry: Its relation with ground-pecking

Abstract It was shown that the motivation for non-aggressive pecking at conspecifics varies along with ground-pecking motivation. This supported the view of a common regulating mechanism. It was concluded that feather-pecking is to be considered as redirected ground-pecking, and hence that the latter is an important parameter in experiments comparing the risk of different environmental factors concerning the development of feather-pecking.

Effects of environmental enrichment on behavioral responses to novelty, learning, and memory, and the circadian rhythm in cortisol in growing pigs

Previously we showed that pigs reared in an enriched environment had higher baseline salivary cortisol concentrations during the light period than pigs reared under barren conditions. In the present experiment, it was investigated whether these higher baseline salivary cortisol concentrations were a real difference in cortisol concentration or merely represented a phase difference in circadian rhythm. The effects of different cortisol concentrations on the behavioral responses to novelty and learning and long-term memory in a maze test were also studied in enriched and barren housed pigs. At 9 weeks of age enriched and barren housed pigs did not differ in baseline salivary cortisol concentrations nor in circadian rhythm, but at 22 weeks of age barren housed pigs had a blunted circadian rhythm in salivary cortisol as compared to enriched housed pigs. The differences in baseline salivary cortisol concentrations between enriched- and barren-housed pigs are age-dependent, and become visible after 15 weeks of age. Enriched- and barren-housed piglets did not differ in time spent on exploration in the novel environment test. Barren-housed pigs had an impaired long-term memory in the maze test compared to enriched-housed pigs; however, no differences in learning abilities between enriched- and barren-housed pigs were found. Because blunted circadian cortisol rhythms are often recorded during states of chronic stress in pigs and rats or during depression in humans, it is suggested that the blunted circadian rhythm in cortisol in barren-housed pigs similarily may reflect decreased welfare.

Quantifying aspects of young horses’ temperament: consistency of behavioural variables

Performance of horses, whether in sports or in leisure, depends on both physical abilities as well as temperament. The aim of the present work was to measure individual variation and consistency of behavioural variables, related to temperament, in young horses of the same breed and age, and reared under controlled housing conditions and management. A total of 41 Dutch Warmblood horses were tested at 9, 10, 21 and 22 months of age in two behavioural tests, i.e. the novel object test and the handling test. In the novel object test horses were confronted with an open umbrella that was lowered from the ceiling. In the handling test horses were led by a human to cross a bridge. Per test, behavioural variables in the following behavioural classes were observed: locomotor activity, latency times, postural expressions and vocalisations. Within years, all behavioural variables in the handling test, and all but two in the novel object test were positively correlated (0.36<Rs<0.81, P<0.05). For both tests, at 9, 10, 21 and 22 months of age, a principal component analysis (PCA) was carried out to examine whether there were indications for underlying components of these individual behavioural variables that could possibly serve as measures for temperamental traits. The first component in the novel object test could be regarded as ‘flightiness’ and the second as ‘sensitiveness’. In the handling test, the first component was suggested to relate to ‘patience’, the second component to ‘willingness to perform’. The temperamental trait ‘flightiness’ (novel object test) as well as the temperamental trait ‘patience’ (handling test) were positively correlated within both years (0.36<Rs<0.65, P<0.05). For the traits ‘sensitiveness’ (novel object test) and ‘willingness to perform’ (handling test) a positive correlation was only found within the first year (0.44<Rs<0.57, P<0.01). A few individual behavioural variables showed consistency over years. Additionally, just one out of four temperamental traits, namely ‘flightiness’, proved to be consistent over years (Rs=0.49, P<0.01). The temperamental trait ‘patience’ showed a trend between years (Rs=0.31, 0.05<P<0.1). It is concluded that the behavioural tests employed in the present study can be used to reliably identify individual behavioural variables and temperamental traits in young horses. Long-term consistency, i.e. between subsequent years, could not be demonstrated convincingly. Nevertheless, future work may indicate that employing the same approach and considering an even longer time period or different phases of the horse’s life, long-term consistency does exist.

Personalities in female domesticated pigs: behavioural and physiological indications

The inconclusive evidence so far on the existence of distinct personality types in domesticated pigs, led us to perform the present experiment. A total of 128 gilts from 31 sows were systematically studied from birth to slaughter in two identical trials. Intra-test consistency in individual behavioural and/or physiological reactions was studied in three different tests. We were not able to show consistencies in reactions of gilts over time to a backtest (at 2-4 days and 4 weeks of age) and to a novel environment test (at 10 and 24 weeks of age). Individual aggression, however, as measured in a group-feeding competition test in stable groups (at 10 and 24 weeks of age), proved to be highly consistent. Explanations for these discrepancies in intra-test consistencies are critically discussed. Inter-test consistencies were determined by relating the individual reactions of gilts to the backtest to various characteristics and responses to tests at a later age. The highest correlations were found when resistance in the first backtest was involved. No evidence was found for the existence of specific isolated categories of animals with respect to this resistance. For further analysis, extreme responding gilts in the first backtest (roughly the top and bottom 25% of the distribution) were classified as low resistant (LR; 4 escape attempts; n = 45). By comparisons of mean responses of LR and HR gilts within groups, we have established a relationship between the backtest and several other variables. Behaviourally, the HR gilts showed more aggression in the group-feeding competition tests. Also, in the competition for the most productive teats at the anterior, a predominant position of HR piglets at this site was observed during the suckling period. The latter piglets also gained more weight during this period than LR ones. Compared to HR pigs, in the first novel environment test LR pigs hesitated longer to leave their home pens and to contact a human, but no difference in their locomotory behaviour was observed. Contrasts between LR and HR pigs in the second novel environment test were reduced or absent. Physiologically, when compared to HR gilts, LR ones had a higher reactivity of the hypothalamic-pituitary-adrenocortical (HPA) system. This was shown by higher cortisol responses to the first novel environment test, to routine weighing at 25 weeks of age, and to administration of a high dose of ACTH. It is discussed that these findings for LR and HR gilts, may provide support for the existence of behavioural and physiological responses in pigs, resembling those of proactive and reactive rodents

The Circadian Rhythm of Salivary Cortisol in Growing Pigs: Effects of Age, Gender, and Stress

This experiment was designed to examine circadian rhythmicity of cortisol in saliva of growing pigs, in relation to age, gender, and (time of) stressor application. Additionally, the acute cortisol response to a stressor was studied. Five groups, each consisting of 3 barrows and 3 gilts, were involved in the experiment. In a Control Group, saliva samples were taken at 1-h intervals at 12, 16, 20, and 24 weeks of age. Within 1 week, rhythmicity of cortisol was assessed during two 24-h spans (Monday and Friday). Rhythm characteristics were evaluated by cosinor analysis, describing the rhythm by several parameters. In 2 groups at 12 weeks and 2 other groups at 20 weeks of age, a stressor was applied (4 h of isolation) on Thursday morning or evening. Again, rhythmicity was assessed on Monday and Friday by sampling at 2-h intervals. Acute cortisol effects were studied by sampling at several time-points during isolation. Between 12 and 24 weeks of age, basal cortisol concentrations decreased and a rather stable and adult circadian rhythm was reached at 20 weeks of age. Average basal cortisol concentrations were higher in barrows than in gilts. Furthermore, after isolation, the amplitude of the rhythm was increased in barrows but was unchanged in gilts. The rhythm was more unstable and the maximum value tended to shift only after evening isolation. Stressor timing, but also age, was found to affect average cortisol concentrations. Moreover, stressor timing was important for the acute cortisol response: the increase was higher in the morning. The results of this study emphasize the importance of considering the circadian rhythmicity of cortisol, in relation to age, gender, and (time of) stressor application, when studying the cortisol response of animals to stressors.

Plasma catecholamine and corticosterone levels during manual restraint in chicks from a high and low feather pecking line of laying hens

Neurosympathetic and adrenal responsivity to manual restraint was studied in two White Leghorn chicken lines which differ in their tendency to feather peck. Blood samples were taken from freely moving cannulated birds during resting conditions and during manual restraint (placing the bird on its side for 8 min) to determine plasma noradrenaline. adrenaline and corticosterone. Plasma corticosterone levels in low feather peckers (LFP) were significantly higher during resting conditions and restraint as compared to the high feather peckers (HFP). In response to manual restraint the HFP hens showed a significantly larger plasma noradrenaline response than LFP hens. No difference in plasma adrenaline levels was observed under resting conditions or during manual restraint.

Some observations on the development of feather-pecking in poultry

Abstract In 2 experiments, observations were made on the development of feather-pecking in poultry. In the first experiment, 4 groups of 8 pullets each were observed from hatching until 17 weeks of age. Two groups were housed on litter (L-groups), two groups on slatted floors (NL-groups). Pecking at conspecifics increased in NL-groups with time, whereas it tended to decrease in L-groups. Moreover, in the latter, about 20% of the pecks was directed to particles on the plumage of other birds and about 25% to feathers. In the NL-groups, these percentages were about 1 and 55, respectively. In experiment 2, half of the animals from each floor-type was transferred to the other type of flooring material. Most striking was that in the group of animals reared on litter and changed to slats, feather-pecking increased significantly with the duration of the experiment. In both experiments, floor-pecking was high in groups with a low frequency of pecking at conspecifics and vice versa. It is concluded that the results strongly support the view that feather-pecking evolves as “misdirected” food-pecking.