Linda J. Keeling

Assessment of positive emotions in animals to improve their welfare

It is now widely accepted that good welfare is not simply the absence of negative experiences, but rather is primarily the presence of positive experiences such as pleasure. However scientific investigation of positive emotions has long been neglected. This paper addresses two main issues: first, it reviews the current state of scientific knowledge that supports the existence of positive affective states in animals and, second, it suggests possible applications of this knowledge that may enhance quality of life under animal management conditions. In the first part of the paper, recent advances in psychology and neuroscience are reviewed to provide pragmatic frameworks based on cognitive processes (such as positive anticipation, contrast and controllability) for further investigations of positive emotions in animals. Thereafter, the neurobiological bases of positive emotions are highlighted in order to identify behavioral and physiological expressions of positive experiences in animals. Monitoring both the autonomic nervous system (via heart rate and its variability) and the immune system could offer relevant tools to better assess emotional states in animals, complementary to classical adrenocortical measures. In the second part of the paper, useful strategies for enhancing positive experiences (such as physical, social and cognitive enrichment or putative genetic selection) are outlined. Then this paper emphasizes practical applications for assessing and promoting positive emotions that may help in providing animals with a better quality of life. Play, affiliative behaviors and some vocalizations appear to be the most promising convenient indicators for assessing positive experiences in laboratory and farm animals under commercial conditions.

Effect of rearing factors on the prevalence of floor eggs, cloacal cannibalism and feather pecking in commercial flocks of loose housed laying hens

1. Effects of rearing conditions on behavioural problems were investigated in a cohort study of commercial flocks of laying hens housed in 2 different loose housing systems. The sample population was 120 385 laying hens from 59 flocks of various hybrids at 21 different farms. 2. Logistic regression modelling was used to test the effects of selected factors on floor eggs, cloacal cannibalism and feather pecking. In addition to early access to perches or litter, models included hybrid, stocking density, group size, housing system, age at delivery, identical housing system at the rearing farm and at the production farm and, in models for floor eggs and cloacal cannibalism, nest area per hen. Odds ratios were calculated from the results of the models to allow risk assessment. 3. No significant correlations were found between the prevalence of floor eggs, cloacal cannibalism and feather pecking. 4. Access to perches from not later than the 4th week of age decreased the prevalence of floor eggs during the period from start-of-lay until 35 weeks of age, odds ratio 0-30 (P<0-001). Furthermore, early access to perches decreased the prevalence of cloacal cannibalism during the production period, odds ratio 0-46 (P=0.03). 5. No other factor had a significant effect in these models. Although it was not significant, early access to litter had a non-significant tendency to reduce the prevalence of feather pecking.

Relationship between feather pecking and ground pecking in laying hens and the effect of group size

The aim of this experiment was to study the relationship between feather pecking and ground pecking in laying hens and the effect of group size on feather pecking behaviour. Hisex White hens were kept in floor pens in group sizes of 15, 30, 60 and 120 birds, each with four replicates. Behavioural observations were performed at four different ages and focused on the number of feather pecks and aggressive pecks, both given and received. The part of the body pecked and the location of the bird was recorded as well as the number of pecks made to the floor, feeder and drinker.The results showed that most feather pecking activity occurred in the largest group size (120 birds) and there was some evidence of an increasing frequency of aggressive pecks with increasing group size. The parts of the body which were targets for feather pecking varied depending on the location of the bird giving the peck and the bird receiving it. When looking at the behaviour of individuals, birds doing a lot of feather pecking also showed more ground pecking.

Effect of manipulating feathers of laying hens on the incidence of feather pecking and cannibalism

Feather pecking is a problem in commercial laying hens, particularly in loose-housing systems, where many hens can be affected by only a few feather peckers. In addition, feather pecking can become an even larger problem if it spreads throughout the flock. There are several possible ways that feather pecking may spread. The simplest way is that one hen may damage the feathers of a hen, and another hen may find the damaged feathers an attractive pecking target. The aim of this experiment was to determine if damaged feathers were feather-pecked more than undamaged feathers on the same body area, and to determine whether some types of feather-body area manipulations were preferred over others as pecking stimuli. Manipulations involved damaging the feathers on the rump, tail or belly of different hens, with two or three levels of severity of manipulation at each body area. Sixteen groups of 11 Lohmann Brown hens between 26 and 28 weeks were observed with the recipient, the feather pecker and the body area that was pecked all being recorded. The feather pecks were classified separately as either gentle or severe. Damaged feathers received significantly more severe feather pecks than undamaged feathers. There were also more gentle feather pecks to damaged feathers, although this did not reach statistical significance. The feather-body area manipulations that received the greatest number of severe feather pecks were the tail feathers when they were cut very short, the rump feathers when they were trimmed, and the rump when feathers were removed. These results support the suggestion that feather pecking does indeed spread through flocks by damaged feathers becoming an attractive target for feather-pecking behaviour. An unexpected result of performing the feather manipulations was an outbreak of cannibalism in half of the experimental groups. Even though there was no visible damage to the skin of the hens after having the feathers manipulated, 13 of the 16 attacked hens were wounded on the part of the body where the feathers had been damaged in some way.

Stocking density effects on broiler welfare: Identifying sensitive ranges for different indicators

Although stocking density is perceived as a topic of major importance, no consensus has been reached on what density would allow for good welfare. In the present study, the welfare of 4 replicates of birds stocked at 8, 19, 29, 40, 45, 51, 61, and 72 broilers per pen (or 6, 15, 23, 33, 35, 41, 47, and 56 kg actually achieved BW/m(2)) was studied using 6 welfare indicators. Density did not affect bursa weight, mortality, or concentrations of corticosterone metabolites in droppings but did influence leg health (P = 0.015) and footpad and hock dermatitis (P < 0.001) and tended to influence fearfulness (P = 0.078). However, not every increase in density or group size, or both, led to poorer welfare for the affected indicators: leg health and fearfulness showed unexpected peaks at intermediate densities. Furthermore, the indicators were influenced at different densities: leg strength showed a steep decrease from 6 to 23 kg/m(2), hock dermatitis rose from 35 to 56 kg/m(2), and footpad dermatitis and fearfulness were only significantly higher at the highest density of 56 kg/m(2). No threshold stocking density above which all aspects of welfare were suddenly altered was found in this study. Instead, different aspects of welfare were influenced at different densities or group sizes, or both. Thus, evaluating the effects of stocking density on welfare as a whole would require either identification of acceptable levels for each separate indicator or a weighting of the indicators in an integrated welfare score. A tentative attempt to such an integration, made using equal weights for all parameters, showed a decrease in welfare as density increased (P < 0.001). The lowest 2 densities (6 and 15 kg/m(2)) scored better than most middle densities (23, 33, 35, and 47 kg/m(2)), whereas all densities scored better than the highest density (56 kg/m(2)).

Chicken genomics: Feather-pecking and victim pigmentation

Feather-pecking in domestic birds is associated with cannibalism and severe welfare problems. It is a dramatic example of a spiteful behaviour in which the victims fitness is reduced for no immediate direct benefit to the perpetrator and its evolution is unexplained. Here we show that the plumage pigmentation of a chicken may predispose it to become a victim: birds suffer more drastic feather-pecking when the colour of their plumage is due to the expression of a wild recessive allele at PMEL17, a gene that controls plumage melanization, and when these birds are relatively common in a flock. These findings, obtained using an intercross between a domestic fowl and its wild ancestor, have implications for the welfare of domestic species and offer insight into the genetic changes associated with the evolution of feather-pecking during the early stages of domestication.

Feather pecking in chickens is genetically related to behavioural and developmental traits

Feather pecking (FP) is a detrimental behaviour in chickens, which is performed by only some individuals in a flock. FP was studied in 54 red junglefowl (ancestor of domestic chickens), 36 White Leghorn laying hens, and 762 birds from an F(2)-intercross between these two lines. From all F(2)-birds, growth and feed consumption were measured. Age at sexual maturity and egg production in females, and corticosterone levels in males were also measured. From 333 F(2)-birds of both sexes, and 20 parental birds, body composition with respect to bone mineral content, muscle and fat was obtained by post-mortem examinations using Dual X-Ray Absorptiometry (DXA). In femurs of the same birds, the bone density and structure were analysed using DXA and Peripheral Quantitative Computerized Tomography (pQCT), and a biomechanical analysis of bone strength was performed. Furthermore, plumage condition was determined in all birds as a measure of being exposed to feather pecking. Using 105 DNA-markers in all F(2)-birds, a genome-wide scan for Quantitative Trait Loci (QTL), associated with the behaviour in the F(2)-generation was performed. FP was at least as frequent in the red junglefowl as in the White Leghorn strain studied here, and significantly more common among females both in the parental strains and in the F(2)-generation. In the F(2)-birds, FP was phenotypically linked to early sexual maturation, fast growth, weak bones, and, in males, also high fat accumulation, indicating that feather peckers have a different resource allocation pattern. Behaviourally, F(2) feather peckers were more active in an open field test, in a novel food/novel object test, and in a restraint test, indicating that feather pecking might be genetically linked to a proactive coping strategy. Only one suggestive QTL with a low explanatory value was found on chromosome 3, showing that many genes, each with a small effect, are probably involved in the causation of feather pecking. There were significant effects of sire and dam on the risk of being a victim of feather pecking, and victims grew faster pre- and post-hatching, had lower corticosterone levels and were less active in a restraint test. Hence, a wide array of behavioural and developmental traits were genetically linked to FP.

Night-time roosting in laying hens and the effect of thwarting access to perches.

Free-living hens roost on branches in trees at night, and laying hens in aviary systems or cages provided with perches also make extensive use of these for night-time roosting. It is therefore suggested that roosting on perches is important to the hens and that domestic hens should be provided with perches in order to promote welfare. However, no study has addressed the question of motivation for roosting. In the present experiment, we studied undisturbed roosting behaviour and the reaction of commercial laying hens when roosting on perches was thwarted. Fifty-two adult hens (Lohmann Selected Leghorn, LSL) were kept in two groups of 26 hens in litter pens with perches at heights of 23, 43 and 63 cm. Behaviour was observed for 60 min starting at lights-off, registering the number of hens on each perch level. The hens started to get onto the perch immediately and within 10 min after lights-off, more than 90% of the hens were on the perch. All hens roosted close together on the top perch. In a second experiment, 24 hens were kept in eight groups of three birds each in experimental pens equipped with perches. Birds were tested in four different situations: (1) the pen unchanged (Base), (2) the perch covered with plexiglass (PCov), (3) the perch removed (PRem) and (4) the unchanged pen (Post). The order of PCov and PRem alternated between groups in a balanced manner and all groups of birds experienced all four treatments. The hens were observed for 60 min from lights-off using focal sampling. For comparisons, the Post treatment served as the control. In the treatments where perching was not possible, the hens spent less time sitting (p=0.042), and also tended to spend more time standing (p=0.06), than in the control. Furthermore, the hens moved more (p=0.042) when the perch was inaccessible, and when the perch was visible but inaccessible they also showed more attempts to take off (p=0.042). These findings can be interpreted as increased frustration and/or exploration, probably to find an alternative roosting site. Together with the high use of perches for night-time roosting under undisturbed conditions, these results indicate that laying hens are motivated to perch and imply that hens kept under conditions where perching is not possible may experience reduced welfare.

Group size and perching behaviour in young domestic fowl.

To test the hypothesis that young domestic fowl perform less perch-related antipredator behaviour with increasing group size, White Leghorn pullets were reared in four replicate groups of 15, 30, 60 and 120 at a constant density of 5 birds/m(2). Each pen contained perches 20, 40 and 60cm above the ground. Perch space per bird per perch level was the same for all groups. It was predicted that, with increasing group size, domestic fowl would (1) spend less time on perches (i.e. more time down on the floor); (2) be less vigilant while perching; (3) spend relatively more time preening down on the floor. As predicted, the proportion of 3- to 18-week-old birds roosting on perches during scans throughout the photoperiod decreased with increasing group size, from 41+/-1.7% in groups of 15 birds to 33+/-1.6% in groups of 120 birds. This effect was due to reduced use of the lower perches; use of the highest perches was high at all group sizes. The proportion of birds vigilant on the highest perches of those present on that perch level decreased with increasing group size. The proportion of birds engaged in the vulnerable activity of preening down on the floor increased with group size. The frequency of transitions between floor and perches was not affected by group size but birds received more disturbances from other birds when on the top perch level in the larger groups. Thus, the decline in vigilance on the top perch level with increasing group size was not due to reduced disturbance from other birds. In conclusion, despite domestication and protection from non-human predators, changes in the use of perches by young domestic fowl with increasing group size were consistent with the antipredator hypothesis.

Dynamics of aggression in the domestic fowl

Abstract We conducted three experiments designed to investigate the effect of group size (GS) on the frequencies of different types of aggressive acts performed by 11–21-week-old White Leghorn pullets. In these experiments, we systematically controlled for bird density, pen size, and previous social experience in groups of different sizes. Aggression was measured in the context of competition for a limited supply of highly attractive food provided in two patches (central and peripheral) in addition to the regular (ad libitum) food supply. The birds were observed in their original groups of 15, 30, 60 and 120 birds (3–4 replicate groups per GS), reared together since hatch, or in temporary sub-groups of the original groups. In the first experiment (space per bird constant, pen size and GS history varying), the number of aggressive pecks delivered per food patch declined as GS increased from 15 to 120 birds. In the second experiment, utilising birds from the groups of 120 only (space per bird varying, pen size and GS history constant), the number of aggressive pecks delivered per patch increased as temporary GS increased from 15 to 60 birds. In the third experiment (space per bird and pen size constant, GS history varying), the number of aggressive pecks delivered per patch was constant among groups of 15 birds obtained from the groups of 15, 30, 60 and 120 birds. In all experiments, the central food patch was exploited sooner than the peripheral one. The actual number of birds at a food patch at a given time was more important in determining the frequency of aggression at the food patch than the number of birds present in an enclosure or previous GS experience. When a high number of individuals was present at a food patch, aggression at that patch was relatively infrequent. The results support the hypothesis that aggression in the domestic fowl is a dynamic process, with decisions about aggressive behaviour being made facultatively according to the relative costs and benefits of different behavioural strategies at a given time and place rather than being fixed obligatorily at a constant level according to the number of birds present in an enclosure and their ability to form dominance relationships.