Prenatal and Early Postnatal Behavioural Programming in Laying Hens, With Possible Implications for the Development of Injurious Pecking.

Injurious pecking (IP) represents a serious concern for the welfare of laying hens (Gallus gallus domesticus). The risk of IP among hens with intact beaks in cage-free housing prompts a need for solutions based on an understanding of underlying mechanisms. In this review, we explore how behavioural programming via prenatal and early postnatal environmental conditions could influence the development of IP in laying hens. The possible roles of early life adversity and mismatch between early life programming and subsequent environmental conditions are considered. We review the role of maternal stress, egg conditions, incubation settings (temperature, light, sound, odour) and chick brooding conditions on behavioural programming that could be linked to IP. Brain and behavioural development can be programmed by prenatal and postnatal environmental conditions, which if suboptimal could lead to a tendency to develop IP later in life, as we illustrate with a Jenga tower that could fall over if not built solidly. If so, steps taken to optimise the environmental conditions of previous generations and incubation conditions, reduce stress around hatching, and guide the early learning of chicks will aid in prevention of IP in commercial laying hen flocks.

Examining affective structure in chickens: valence, intensity, persistence and generalization measured using a Conditioned Place Preference Test.

When measuring animals' valenced behavioural responses to stimuli, the Conditioned Place Preference (CPP) test goes a step further than many approach-based and avoidance-based tests by establishing whether a learned preference for, or aversion to, the location in which the stimulus was encountered can be generated. We designed a novel, four-chambered CPP test to extend the capability of the usual CPP paradigm to provide information on four key features of animals' affective responses: valence, scale, persistence and generalization. Using this test, we investigated the affective responses of domestic chickens (Gallus gallus domesticus) to four potentially aversive stimuli: 1. Puffs of air; 2. Sight of (robotic) snake; 3. Sprays of water; 4. Sound of conspecific alarm calls. We found conditioned avoidance of locations associated with the air puffs and water sprays (Friedman's χ2 (3) = 13.323 p > .005; χ2 (3) = 14.235 p > .005), but not with the snake and alarm calls. The scale of the learned avoidance was similar for the air puff and water spray stimuli, but persistence and generalization differed. We conclude that the four chambered CPP test can have a valuable role to play in making multi-feature measurements of stimulus-generated affective responses, and we highlight the value of such measurements for improving our understanding of the structure of affect in chickens and other animals.

Socially-mediated arousal and contagion within domestic chick broods.

Emotional contagion - an underpinning valenced feature of empathy - is made up of simpler, potentially dissociable social processes which can include socially-mediated arousal and behavioural/physiological contagion. Previous studies of emotional contagion have often conflated these processes rather than examining their independent contribution to empathic response. We measured socially-mediated arousal and contagion in 9-week old domestic chicks (n = 19 broods), who were unrelated but raised together from hatching. Pairs of observer chicks were exposed to two conditions in a counterbalanced order: air puff to conspecifics (AP) (during which an air puff was applied to three conspecifics at 30 s intervals) and control with noise of air puff (C) (during which the air puff was directed away from the apparatus at 30 s intervals). Behaviour and surface eye temperature of subjects and observers were measured throughout a 10-min pre-treatment and 10-min treatment period. Subjects and observers responded to AP with increased freezing, and reduced preening and ground pecking. Subjects and observers also showed reduced surface eye temperature - indicative of stress-induced hyperthermia. Subject-Observer behaviour was highly correlated within broods during both C and AP conditions, but with higher overall synchrony during AP. We demonstrate the co-occurrence of socially-mediated behavioural and physiological arousal and contagion; component features of emotional contagion.

Behavioural and physiological responses of laying hens to automated monitoring equipment.

Automated monitoring of behaviour can offer a wealth of information in circumstances where observing behaviour is difficult or time consuming. However, this often requires attaching monitoring devices to the animal which can alter behaviour, potentially invalidating any data collected. Birds often show increased preening and energy expenditure when wearing devices and, especially in laying hens, there is a risk that individuals wearing devices will attract aggression from conspecifics. We studied the behavioural and physiological response of 20 laying hens to backpacks containing monitoring devices fastened with elastic loops around the wing base. We hypothesised that backpacks would lead to a stress-induced decrease in peripheral temperature, increased preening, more aggression from conspecifics, and reduced bodyweights. This was evaluated by thermography of the eye and comb (when isolated after fitting backpacks), direct observations of behaviour (when isolated, when placed back into the group, and on later days), and weighing (before and after each 7-day experimental period). Each hen wore a backpack during one of the two experimental periods only and was used as her own control. Contrary to our hypothesis, eye temperature was higher when hens wore a backpack (No backpack: 30.2 °C (IQR: 29.0-30.6) vs. Backpack: 30.9 °C (IQR: 30.0-32.0), P < 0.001). Eye temperature of hens wearing a backpack was strongly correlated to the time spent preening (rs = 0.8, P < 0.001), suggesting that the higher temperatures may have been due to preening itself, or to a low head position or decreased heat dissipation when preening under the wings. Aggressive behaviour was very rare and no effect of the backpacks was found. In line with our hypothesis, backpacks increased preening on the day of fitting, both when isolated (No backpack: 0% (IQR: 0-1) vs. Backpack: 22% (IQR: 1-43), P < 0.01) and when back in the group (No backpack: 0% (IQR: 0-27) vs. Backpack: 43% (IQR: 5-77), P < 0.001). However, no effect on preening was observed 2-7 days afterwards. Other behavioural changes suggested that on the day of fitting hens prioritized attempts to (re)move the backpack and were less attentive to their surroundings. However, only equipment pecking (i.e., pecking the backpack or leg rings) was still affected 2-7 days after fitting (No backpack: 0 pecks/hen/minute (IQR: 0-0), vs. Backpack: 0 (IQR: 0-0.07), P < 0.05). We found no effect of our backpacks on bodyweight. In conclusion, our backpacks seem suitable to attach monitoring equipment to hens with only a very minor effect on their behaviour after a short acclimation period (≤2 days).

Influences of Maternal Care on Chicken Welfare.

In domestic chickens, the provision of maternal care strongly influences the behavioural development of chicks. Mother hens play an important role in directing their chicks' behaviour and are able to buffer their chicks' response to stressors. Chicks imprint upon their mother, who is key in directing the chicks' behaviour and in allowing them to develop food preferences. Chicks reared by a mother hen are less fearful and show higher levels of behavioural synchronisation than chicks reared artificially. In a commercial setting, more fearful chicks with unsynchronised behaviour are more likely to develop behavioural problems, such as feather pecking. As well as being an inherent welfare problem, fear can also lead to panic responses, smothering, and fractured bones. Despite the beneficial effects of brooding, it is not commercially viable to allow natural brooding on farms and so chicks are hatched in large incubators and reared artificially, without a mother hen. In this review we cover the literature demonstrating the important features of maternal care in domestic chickens, the behavioural consequences of deprivation and the welfare implications on commercial farms. We finish by suggesting ways to use research in natural maternal care to improve commercial chick rearing practice.

Towards a 'Good Life' for Farm Animals: Development of a Resource Tier Framework to Achieve Positive Welfare for Laying Hens.

The concept of a 'good life' recognises the distinction that an animal's quality of life is beyond that of a 'life worth living', representing a standard of welfare substantially higher than the legal minimum (FAWC, 2009). We propose that the opportunities required for a 'good life' could be used to structure resource tiers that lead to positive welfare and are compatible with higher welfare farm assurance schemes. Published evidence and expert opinion was used to define three tiers of resource provision (Welfare +, Welfare ++ and Welfare +++) above those stipulated in UK legislation and codes of practice, which should lead to positive welfare outcomes. In this paper we describe the principles underpinning the framework and the process of developing the resource tiers for laying hens. In doing so, we summarise expert opinion on resources required to achieve a 'good life' in laying hens and discuss the philosophical and practical challenges of developing the framework. We present the results of a pilot study to establish the validity, reliability and feasibility of the draft laying hen tiers on laying hen production systems. Finally, we propose a generic welfare assessment framework for farm animals and suggest directions for implementation, alongside outcome parameters, that can help define and promote a future 'good life' for farm animals.

No evidence for emotional empathy in chickens observing familiar adult conspecifics.

The capacity of animals to empathise is of high potential relevance to the welfare of group-housed domestic animals. Emotional empathy is a multifaceted and multilayered phenomenon which ranges from relatively simple processes such as emotional matching behaviour to more complex processes involving interaction between emotional and cognitive perspective taking systems. Our previous research has demonstrated that hens show clear behavioural and physiological responses to the mild distress of their chicks. To investigate whether this capacity exists outside the mother/offspring bond, we conducted a similar experiment in which domestic hens were exposed to the mild distress of unrelated, but familiar adult conspecifics. Each observer hen was exposed to two replicates of four conditions, in counterbalanced order; control (C); control with noise of air puff (CN); air puff to conspecific hen (APC); air puff to observer hen (APH). During each test, the observer hens' behaviour and physiology were measured throughout a 10 min pre-treatment and a 10 min treatment period. Despite showing signs of distress in response to an aversive stimulus directed at themselves (APH), and using methodology sufficiently sensitive to detect empathy-like responses previously, observer hens showed no behavioural or physiological responses to the mild distress of a familiar adult conspecific. The lack of behavioural and physiological response indicates that hens show no basis for emotional empathy in this context.

Consistency, transitivity and inter-relationships between measures of choice in environmental preference tests with chickens.

Choice experiments are a widespread method of assessing the welfare requirements of domestic animals but prior predictions about what animals should choose to maximise their welfare are not possible. It is therefore important to analyse the data from sets of choice tests to establish whether intra-set and inter-set consistency in choice occurs. Strong inter-set consistency (transitivity) would provide support for the assumption that animals are maximising a complex utility (welfare) that may map onto their subjective state. We housed 56 hens Gallus gallus sequentially in three environments, experienced as three sets (A vs B; B vs C; A vs C), over a 40-week period. At the end of each set, each chicken made six choices between the items in that set, using T-maze testing procedures. The chickens showed significantly greater within-set consistency than expected by chance (p<0.001 for all sets). Although different chickens exhibited different environmental preferences, we also found that a high within-set consistency was associated with faster decision-making (p<0.001). When the three sets of choice trials for each hen were compared there was some evidence that the birds made more transitive choices (indicating consistent choices between sets) than expected by chance.

Towards humane end points: behavioural changes precede clinical signs of disease in a Huntington's disease model.

The number of animals used in science is increasing, bringing a concomitant obligation to minimize suffering. For animals with progressive conditions, euthanasia at a 'humane end point' is advised if the end point is scientifically valid, predictive and accurate. Our aim was to test the hypothesis that behavioural changes would reliably precede clinical signs of disease in a progressive neurological model, using retrospective analysis. We observed 100 pair-housed female R6/1 transgenic Huntington's disease (HD) mice and 28 pair-housed female wild-type (WT) mice in standard- or resource-enriched cages. Disease progression was monitored until one member of each HD pair reached a pre-defined end point based on pathological symptoms (HD end). This mouse was then euthanized together with its cage mate (HD other) and any matched WT pairs. At euthanasia, HD mice had significantly greater absolute and relative organ weights, and significantly higher alpha1 acid glycoprotein concentrations than WT mice, indicating reduced welfare. HD mice initially showed significantly greater use of cage resources than WT mice but this declined progressively. Steeper declines, and earlier cessation, in the use of some climbing and exploration resources occurred in the HD end mice compared with the HD other mice. Behavioural change can be an early indicator of disease onset.