Behavioural and physiological stress responses to handling in wild guanacos

Both ethology and physiology can provide insights about stress conditions in wild animals during handling. Lama guanicoe (guanaco) is the major large herbivore inhabiting arid zones in Argentina. Since 2005, the Payún Matrú Cooperative, a business established and run by native people, has captured and handled guanacos for shearing at Payunia Reserve, Mendoza Province, Argentina, to market the species’ fibre for clothing and other products. The aim of our study was to evaluate the behavioural and physiological stress responses to handling of wild guanacos during these shearing activities. We evaluated stress conditions in guanacos by monitoring the frequency of agonistic behaviour in the holding corrals, following capture, the rate of vocalisation and agonistic behaviour during handling, and by estimating cortisol levels in handled animals. Crowding increased agonistic behaviour; this appeared to be related to dominance conflicts between males in the corrals. Time of corralling increased rate of agonistic behaviour and vocalisations during handling. The rate of behavioural stress responses was negatively related to cortisol levels in males; which may be due to animal fatigue or because behavioural stress responses may help mitigate physiological stress responses. Incorporating our results into animal welfare practices could help reduce stress conditions in wild guanacos. Our results suggest that both behavioural and physiological responses are important measures of stress in handled animals, and that behavioural stress responses cannot be used as a simple surrogate for glucocorticoid levels.     

Concurrent modulation of neuronal and behavioural olfactory responses to sex and host plant cues in a male moth

Mating has profound effects on animal physiology and behaviour, not only in females but also in males, which we show here for olfactory responses. In cotton leafworm moths, Spodoptera littoralis, odour-mediated attraction to sex pheromone and plant volatiles are modulated after mating, producing a behavioural response that matches the physiological condition of the male insect. Unmated males are attracted by upwind flight to sex pheromone released by calling females, as well as to volatiles of lilac flowers and green leaves of the host plant cotton, signalling adult food and mating sites, respectively. Mating temporarily abolishes male attraction to females and host plant odour, but does not diminish attraction to flowers. This behavioural modulation is correlated with a response modulation in the olfactory system, as shown by electro-physiological recordings from antennae and by functional imaging of the antennal lobe, using natural odours and synthetic compounds. An effect of mating on the olfactory responses to pheromone and cotton plant volatiles but not to lilac flowers indicates the presence of functionally independent neural circuits within the olfactory system. Our results indicate that these circuits interconnect and weigh perception of social and habitat odour signals to generate appropriate behavioural responses according to mating state.     

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.     

Social structure and indirect genetic effects: genetics of social behaviour

The social environment modulates gene expression, physiology, behaviour and patterns of inheritance. For more than 50 years, this concept has been investigated using approaches that include partitioning the social component out of behavioural heritability estimates, studying maternal effects on offspring, and analysing dominance hierarchies. Recent advances have formalized this ‘social environment effect’ by providing a more nuanced approach to the study of social influences on behaviour while recognizing evolutionary implications. Yet, in most of these formulations, the dynamics of social interactions are not accounted for. Also, the reciprocity between individual behaviour and group‐level interactions has been largely ignored. Consistent with evolutionary theory, the principles of social interaction are conserved across a broad range of taxa. While noting parallels in diverse organisms, this review uses Drosophila melanogaster as a case study to revisit what is known about social interaction paradigms. We highlight the benefits of integrating the history and pattern of interactions among individuals for dissecting molecular mechanisms that underlie social modulation of behaviour.     

Pheromonal control: reconciling physiological mechanism with signalling theory

Pheromones are intraspecific chemical signals. They can have profound effects on the behaviour and/or physiology of the receiver, and it is still common to hear pheromones described as controlling of the behaviour of the receiver. The discussion of pheromonal control arose initially from a close association between hormones and pheromones in the comparative physiological literature, but the concept of a controlling pheromone is at odds with contemporary signal evolution theory, which predicts that a manipulative pheromonal signal negatively affecting the receiver's fitness should not be stable over evolutionary time. Here we discuss the meaning of pheromonal control, and the ecological circumstances by which it might be supported. We argue that in discussing pheromonal control it is important to differentiate between control applied to the effects of a pheromone on a receiver's physiology (proximate control), and control applied to the effects of a pheromone on a receiver's fitness (ultimate control). Critically, a pheromone signal affecting change in the receiver's behaviour or physiology need not necessarily manipulate the fitness of a receiver. In cases where pheromonal signalling does lead to a reduction in the fitness of the receiver, the signalling system would be stable if the pheromone were an honest signal of a social environment that disadvantages the receiver, and the physiological and behavioural changes observed in the receiver were an adaptive response to the new social circumstances communicated by the pheromone.     

Control of behavioural strategies for capricious environments

In addition to seasonal changes in morphology, physiology and behaviour that occur in predictable annual cycles, there are facultative responses to unpredictable events known as labile (i.e. shortlived) perturbation factors. These rapid behavioural and physiological changes have been termed the ‘emergency life-history stage’ and serve to enhance lifetime fitness. There are four major components: (1) proactive/reactive coping styles for responding to psychosocial stress, predation, and so forth; (2) fight-or-flight responses to rapid emergencies such as an attack by a predator or sudden severe storm; (3) ‘take-it-or-leave-it’ behavioural and physiological responses to longer-term perturbations of the physical environment; and (4) sickness behaviour and fever designed to respond to infection. Glucocorticosteroids interact with cytokines and with other hormones in the hypothalamo-pituitary-adrenal cascade and in the autonomic nervous system to initiate and orchestrate the emergency life-history stage within minutes to hours. Some traits of the emergency life-history stage include: redirection of behaviour from a normal life-history stage to increased foraging, irruptive-type migration during the day, enhanced restfulness at night, elevated gluconeogenesis, and recovery once the perturbation passes. These physiological and behavioural changes allow an individual to avoid potential deleterious effects of stress that may result from chronically elevated levels of circulating glucocorticosteroids over days and weeks. Thus, acute rises in glucocorticosteroids following perturbations of the environment may serve primarily as ‘antistress’ hormones, potentially allowing individuals to avoid chronic stress. Several field studies in diverse habitats indicate that individuals in free-living populations show elevated circulating levels of corticosteroids when they are in an emergency life-history stage. Some simple models based on food availability, body condition, social status and life-history stage, may allow predictions of sensitivity of the hypothalamo-pituitary-adrenal axis to labile perturbation factors. Although there is now extensive evidence for behavioural components of the emergency life-history stage in birds, there remains much to be learned about how other vertebrate groups, especially fish, cope with perturbations of the environment. Because of the unpredictable nature of these perturbations, systematic study of behavioural responses to them is not possible and investigators need to be ‘opportunistic’. There is also a growing need to expand our knowledge of these phenomena because human disturbance, global climate change and pollution are all major perturbations of the environment. How vertebrates respond to the unpredictable in general will thus have important conservation value for the future. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Developmental programming: Cumulative effects of increased pre-hatching corticosterone levels and post-hatching unpredictable food availability on physiology and behaviour in adulthood

Prolonged exposure to stress during development can have long-term detrimental effects on health and wellbeing. However, the environmental matching hypothesis proposes that developmental stress programs physiology and behaviour in an adaptive way that can enhance fitness if early environments match those experienced later in life. Most research has focused on the harmful effects that stress during a single period in early life may exert in adulthood. In this study, we tested the potential additive and beneficial effects that stress experienced during both pre- and post-hatching development may have on adult physiology and behaviour. Japanese quail experienced different stress-related treatments across two developmental life stages: pre-hatching corticosterone (CORT) injection, post-hatching unpredictable food availability, both pre- and post-hatching treatments, or control. In adulthood, we determined quails' acute stress response, neophobia and novel environment exploration. The pre-hatching CORT treatment resulted in attenuated physiological responses to an acute stressor, increased activity levels and exploration in a novel environment. Post-hatching unpredictable food availability decreased adults' latency to feed. Furthermore, there were cumulative effects of these treatments across the two developmental stages: quail subjected to both pre- and post-hatching treatments were the most explorative and risk-taking of all treatment groups. Such responses to novel environments could enhance survival in unpredictable environments in later life. Our data also suggest that these behavioural responses may have been mediated by long-term physiological programming of the adrenocortical stress response, creating phenotypes that could exhibit fitness-enhancing behaviours in a changing environment.     

Frequency-dependent physiological trade-offs between competing colour morphs

Evolutionary theory suggests that alternative colour morphs (i.e. genetically controlled phenotypes) may derive similar fitness under frequency-dependent selection. Here we experimentally demonstrate opposing effects of frequency-dependent social environments on plasma hormone levels (testosterone and corticosterone) and immune function between red- and black-headed male morphs of the Gouldian finch (Erythrura gouldiae). Red-headed males are highly sensitive to changes in the social environment, especially towards the relative density of their own aggressive morph, exhibiting high stress responses and immunosuppression in socially competitive environments. In contrast, the non-aggressive black-headed males follow a more passive strategy that appears to buffer them against social stresses. The differential effect of hormones on aggressive behaviour and immune performance reinforces the contrasting behavioural strategies employed by these colour morphs, and highlights the importance of the social environment in determining the individual basis of behavioural and physiological responses.     

Effect of the early social environment on behavioural and genomic responses to a social challenge in a cooperatively breeding vertebrate

The early social environment can have substantial, lifelong effects on vertebrate social behaviour, which can be mediated by developmental plasticity of brain gene expression. Early‐life effects can influence immediate behavioural responses towards later‐life social challenges and can activate different gene expression responses. However, while genomic responses to social challenges have been reported frequently, how developmental experience influences the shape of these genomic reaction norms remains largely unexplored. We tested how manipulating the early social environment of juvenile cooperatively breeding cichlids, Neolamprologus pulcher, affects their behavioural and brain genomic responses when competing over a resource. Juveniles were reared either with or without a breeder pair and a helper. Fish reared with family members behaved more appropriately in the competition than when reared without. We investigated whether the different social rearing environments also affected the genomic responses to the social challenge. A set of candidate genes, coding for hormones and receptors influencing social behaviour, were measured in the telencephalon and hypothalamus. Social environment and social challenge both influenced gene expression of egr‐1 (early growth response 1) and gr1 (glucocorticoid receptor 1) in the telencephalon and of bdnf (brain‐derived neurotrophic factor) in the hypothalamus. A global analysis of the 11 expression patterns in the two brain areas showed that neurogenomic states diverged more strongly between intruder fish and control fish when they had been reared in a natural social setting. Our results show that same molecular pathways may be used differently in response to a social challenge depending on early‐life experiences.     

Social plasticity in fish: integrating mechanisms and function

Social plasticity is a ubiquitous feature of animal behaviour. Animals must adjust the expression of their social behaviour to the nuances of daily social life and to the transitions between life‐history stages, and the ability to do so affects their Darwinian fitness. Here, an integrative framework is proposed for understanding the proximate mechanisms and ultimate consequences of social plasticity. According to this framework, social plasticity is achieved by rewiring or by biochemically switching nodes of the neural network underlying social behaviour in response to perceived social information. Therefore, at the molecular level, it depends on the social regulation of gene expression, so that different brain genomic and epigenetic states correspond to different behavioural responses and the switches between states are orchestrated by signalling pathways that interface the social environment and the genotype. At the evolutionary scale, social plasticity can be seen as an adaptive trait that can be under positive selection when changes in the environment outpace the rate of genetic evolutionary change. In cases when social plasticity is too costly or incomplete, behavioural consistency can emerge by directional selection that recruits gene modules corresponding to favoured behavioural states in that environment. As a result of this integrative approach, how knowledge of the proximate mechanisms underlying social plasticity is crucial to understanding its costs, limits and evolutionary consequences is shown, thereby highlighting the fact that proximate mechanisms contribute to the dynamics of selection. The role of teleosts as a premier model to study social plasticity is also highlighted, given the diversity and plasticity that this group exhibits in terms of social behaviour. Finally, the proposed integrative framework to social plasticity also illustrates how reciprocal causation analysis of biological phenomena (i.e. considering the interaction between proximate factors and evolutionary explanations) can be a more useful approach than the traditional proximate–ultimate dichotomy, according to which evolutionary processes can be understood without knowledge on proximate causes, thereby black‐boxing developmental and physiological mechanisms.     

Under a neighbour's influence: public information affects stress hormones and behaviour of a songbird

Socially acquired information improves the accuracy and efficiency of environmental assessments and can increase fitness. Public information may be especially useful during unpredictable food conditions, or for species that depend on resources made less predictable by human disturbance. However, the physiological mechanisms by which direct foraging assessments and public information are integrated to affect behaviour remain largely unknown. We tested for potential effects of public information on the behavioural and hormonal response to food reduction by manipulating the social environment of captive red crossbills (Loxia curvirostra). Red crossbills are irruptive migrants that are considered sensitive to changes in food availability and use public information in decision making. Here, we show that public information can attenuate or intensify the release of glucocorticoids (i.e. stress hormones) during food shortage in red crossbills. The observed modulation of corticosterone may therefore be a physiological mechanism linking public information, direct environmental assessments and behavioural change. This mechanism would not only allow for public information to affect individual behaviour, but might also facilitate group decision making by bringing group members into more similar physiological states. The results further suggest that stressors affecting entire populations may be magnified in individual physiology through social interactions.     

Low temperature thresholds: are chill coma and CT(min) synonymous?

The effects of sub-lethal low temperatures on insect physiology and behaviour are important determinants of insect activity including foraging, mating, and predation avoidance. A substantial body of research seeks to relate the temperatures at which these activities are compromised to both, climatic conditions at species range limits and underlying physiological processes. The interpretation of this research is complicated by confusion in the names and definition of the responses measured and their associated temperature thresholds. The development of the nomenclature and explanations of the underlying physiological causes are reviewed in order to elucidate the correct sequence of responses/thresholds and associated terminologies. The results of this analysis indicate that: (1) chill coma is a clearly defined, reversible physiological state characterised by the absence of electrophysiological activity. (2) The onset of chill coma begins when low temperatures begin to impair insect behaviour and physiology, and is punctuated by a series of behavioural and/or physiological thresholds or responses. These include the temperatures at which (i) spontaneous movements cease, (ii) coordination is lost to the degree that locomotion becomes impossible, and (iii) chill coma is entered. (3) Confusion has arisen because (a) the term ‘onset of chill coma’ has been used to describe all three of these responses/thresholds and (b) the term CT_min has entered the insect literature from the vertebrate literature. These issues are discussed and a potential solution is proposed to provide clarity and consistency in the future literature.     

Behavioural,endocrine and immune consequences of mixing in weaned piglets

Mixing piglets at weaning increases plasma cortisol concentrations and agonistic behaviour. In contrast to what is observed in older pigs, studies failed to show any effect of social environment on other behavioural variables or on immune function. The lack of effect of mixing may not reflect an absence of stress, but rather the fact that the physiological effects of social reorganisation are masked by the much more important effects of diet change. The aim of this experiment was to evaluate the reactivity of piglets to mixing by dissociating social reorganisation from weaning in itself. For this purpose, the influence of mixing was investigated 5 days after weaning (day 0) in eight control (C) and eight mixed (M) female pigs. Salivary cortisol and behavioural activity were measured from day −1 to day 3. Blood lymphocyte proliferation was measured on days 0 and 3. Cortisol levels were increased after mixing and returned to basal values within 24 h. Blood lymphocyte proliferation was not affected. Mixing increased resting behaviour. Cortisol and behavioural responses were influenced by the social position of individuals in their new group. Piglets seemed to avoid conflicting encounters by diminishing the synchronisation of their activities with their new group. These results suggest that social reorganisation could be stressful for weaned pigs. However, piglets seem to develop behavioural strategies, which could explain the absence of long-term endocrine and immune consequences of mixing.     

The integration of sociality,monoamines and stress neuroendocrinology in fish models: applications in the neurosciences

Animal‐focused research has been crucial for scientific advancement, but rodents are still taking a starring role. Starting as merely supporting evidence found in rodents, the use of fish models has slowly taken a more central role and expanded its overall contributions in areas such as social sciences, evolution, physiology and recently in translational medical research. In the neurosciences, zebrafish Danio rerio have been widely adopted, contributing to our understanding of the genetic control of brain processes and the effects of pharmacological manipulations. However, discussion continues regarding the paradox of function versus structure, when fishes and mammals are compared and on the potentially evolutionarily conserved nature of behaviour across fish species. From a behavioural standpoint, we explore aversive–stress and social behaviour in selected fish models and refer to the extensive contributions of stress and monoaminergic systems. We suggest that, in spite of marked neuroanatomical differences between fishes and mammals, stress and sociality are conserved at the behavioural and molecular levels. We also suggest that stress and sociality are mediated by monoamines in predictable and non‐trivial ways and that monoamines could bridge the relationship between stress and social behaviour. To reconcile the level of divergence with the level of similarity, we need neuroanatomical, pharmacological, behavioural and ecological studies conducted in the laboratory and in nature. These areas need to add to each other to enhance our understanding of fish behaviour and ultimately how this all may lead to better model systems for translational studies.     

Does the environmental background (intensive v. outdoor systems) influence the behaviour of piglets at weaning?

Under intensive pig husbandry, outdoor systems offer a more complex physical and social environment compared with indoor systems (farrowing sheds). As the rearing environment affects behavioural development, it can, therefore, influence behavioural responses of pigs to stressful environments in later stages of production. We tested how the rearing environment influenced behavioural responses to a novel arena test in piglets on the day that they were weaned and mixed into large groups. We recorded video footage and compared the behavioural responses of 30 outdoor-raised and 30 farrowing shed-raised piglets tested in an experimental arena and sequentially exposed to four challenges (each for 5 min) on the day of weaning. Quantitative and qualitative behavioural measures were recorded using time budgets and scoring demeanour or ‘qualitative behavioural expression’ (using Qualitative Behavioural Assessment (QBA)). When held in isolation (challenge 1), both groups were scored as more ‘scared/worried’, while outdoor-raised piglets spent more time eating and jumping against the arena walls. Both groups interacted with a plastic ball (challenge 2: exposure to a novel object) during which they were scored as more ‘playful/curious’ than other challenges. When a food bowl was introduced (challenge 3), farrowing shed-raised piglets were more interested in playing with the food bowl itself, whereas outdoor-raised piglets spent more time eating the feed. Finally, there were no significant differences in social behaviour (challenge 4: introduction of another piglet) between the two groups in terms of the latency to contact each other, amount of time recorded engaged in aggressive/non-aggressive social interactions or QBA scores. Although piglets spent 30% of their time interacting with the other piglet, and half of this time (47%) was engaged in negative interactions (pushing, biting), the levels of aggression were not different between the two groups. Overall, outdoor-raised piglets ate more and were scored as more ‘calm/passive’, whereas farrowing shed-raised piglets spent more time investigating their environment and were scored as more ‘playful/inquisitive’. In conclusion, we did not find differences in behaviour between outdoor-raised and farrowing shed-raised piglets that would highlight welfare issues. The differences found in this study may reflect conflicting affective states, with responses to confinement, neophobia and motivation for exploration evident.     

Effects of husbandry and management systems on physiology and behaviour of farmed and laboratory rabbits

The major issues regarding the welfare of both farmed and laboratory rabbits are reviewed, according to husbandry and management systems. The main stressors that can affect welfare and homeostatic responses in rabbits are also reviewed. An overview of the most widespread housing systems for both farmed and laboratory rabbits is presented. The main problems related to housing and management are identified, in particular those related to individual and group housing, space requirements and group size, as well as human-animal interaction. The effects of psychological and physical stressors on physiology and behaviour are illustrated through examples in various rearing conditions. Psychological stressors include social stress and fear, while physical stressors include environmental variables such as housing system and climatic factors, i.e. heat. Welfare indicators are identified that can be monitored to determine the effects of individual and environmental variables on the animals' possible coping strategies. Physiological indicators include the neuro-endocrine and psycho-neuro-immuno-endocrine measurements, while behavioural indicators include the behavioural repertoire and responses to behavioural tests. Some possible ways to enhance welfare are indicated, such as enrichment of the environment and improved handling procedures.     

Going beyond the limits: effect of clock disruption on human health

Abstract

Synchronisation of organisms’ physiology and behaviour with the external environment is necessary for survival and reproductive fitness. This is critical for human health also. In the past, humans were exposed to predictable natural day and night cycles that allowed the internal clock to synchronise the daily rhythms in physiology and behaviour with the external environment. However, the industrial revolution has made us a 24*7 society and forced the extension of day into night via adoption of artificial light in our lives. This has altered the perception of day and night and made it difficult for the biological processes to synchronise. Such weak synchronisation can be seen in different physiological and behavioural functions that are under circadian control, such as sleep–wake behaviour, melatonin and cortisol rhythms, core body temperature cycle, etc. This also influences the regulatory mechanism at cell and gene levels. Circadian disruption has resulted in increasing incidences of certain cancers, metabolic dysfunction and mood disorders. Several evidence suggest that exposure to aberrant light alters the brain functions that regulate emotion and mood. The present discussion focuses on understanding the effect of circadian disruption on human health, and its various aspects.     

Evidence for activation of endogenous opioid systems in mice following short exposure to stable flies

Biting flies influence both physiology and behaviour of domestic and wild animals. This study demonstrates that brief (30 min) exposure of male and female mice to stable flies leads to significant increases in nociceptive responses, indicative of the induction of analgesia. The biting fly-induced analgesia was mediated by endogenous opioid systems as it was blocked by the prototypic opiate antagonist naloxone. Exposure for 30 min to the bedding of biting fly-exposed mice also induced significant opioid mediated analgesic responses in mice. Exposure to either house flies or the bedding of house fly-exposed mice had no significant effects on nociception. These results indicate that brief exposure to either stable flies, or to olfactory cues associated with mice exposed to stable flies, activates endogenous opioid systems leading to the induction of analgesia and likely other opioid mediated behavioural and physiological stress responses. These results suggest the involvement of endogenous opioid systems in the mediation of the behavioural and physiological consequences of biting fly exposure in domestic and wild animals.     

Social transmission of physiological and behavioural responses to castration in suckling Merino lambs

In social species like sheep, social context can modify both physiological and behavioural responses to stressors and normal behavioural patterns. Presence of conspecifics can ameliorate responses to noxious stimuli, an effect termed social buffering, whereas the presence of a distressed conspecific can invoke a distress response in an observer animal not receiving a noxious organic insult, an effect termed social contagion. Furthermore, synchrony of normal behaviours can occur within a group. Here a range of social contexts were created by grouping suckling lambs undergoing knife, ring or sham castration treatments either homogeneously or heterogeneously by castration treatment in pens within an animal house. The impact of social grouping on cortisol, rectal temperature, pain avoidance behaviours, postural behaviours and synchrony of behaviour in the first 12 h following castration treatment was examined. Irrespective of castration treatment, lambs grouped homogeneously by castration treatment had higher cortisol concentrations across the four time points measured than lambs grouped heterogeneously. They also spent 1.9% more of the time in restless behaviour in the first 1 h following castration, and in the 12 h following castration spent 6.7% more of their time lying ventrally, 4.9% less time standing normally, 1.6% more time walking normally, and 4.9% less time in total standing postures. Interactions between social grouping and castration treatment were not significant for physiological variables, pain related behaviours or postures with the exception of lateral lying which was decreased in knife castrated lambs grouped heterogeneously with other castration treatments from 1.9% (homogeneous grouping) to 0.4% (heterogeneous group). Synchronisation of behaviour was seen for walking in lambs grouped heterogeneously by castration treatment and for feeding at the trough in lambs grouped homogenously by ring and sham castration treatments. Lying and standing respectively, were negatively synchronised (occurred less frequently than predicted by chance alone) in ring and knife castrated lambs grouped homogeneously. The results indicate that mixing lambs that received the three castration treatments within a pen modified the activity profile of the lambs but did not substantially modify the behavioural or physiological changes that are stigmatic of responses to the specific castration treatments. The social contexts contrasted in the study did not result in marked contagion or social buffering between lambs of the measured responses and did not confound comparison of castration treatments. None-the-less, the comparability of responses observed in castration treatment groups penned homogenously or heterogeneously might not necessarily occur in other classes of sheep or in other social contexts. The absence of social transmission of behavioural and physiological responses indicates that grouping non-castrated lambs with castrated lambs did not compromise the welfare of the non-castrated lambs when their mothers were present.