Does audience size influence actors' and spectators' emotions the same way?

Better understanding how audience size influences emotions and behaviours during public performances is of particular importance since it may both impact the level of anxiety and quality of achievement of the performer and alter the degree of appreciation of the observer. We tested this question in a naturalistic setting by analyzing self-assessment questionnaires, Galvanic skin responses and behaviours of actors and spectators during theatrical representations with small, medium and large audiences. We found that: actors and spectators differed in their perception of the effects of audience size; the different components of emotions (cognitive, physiological, behavioural) were affected differently by audience size, which was also modulated by the individual's status; actors and spectators differed in their representation of the others' emotional state. Although our study remains exploratory, our findings highlight the complexity of the audience effect when comparing observers' and performers' emotions.

Could snorts inform us on how horses perceive riding?

Several previous studies have shown that working conditions (including riding) can induce stress in horses. Riders' actions and postures, when inappropriate, induce stress and conflict behaviours during riding and welfare impairment and negative emotional states outside work. Optimistic biases have been found in leisure horses, which, amongst positive management factors, were ridden with loose reins and low hands. Thus, one can wonder whether horses may positively perceive work or parts of it. Indicators of positive emotions are poorly known yet but we recently found that, out of the working context, a non-vocal acoustic signal, snorts, could reflect mild positive emotions in horses. We hypothesized that snorts could help identifying the working phases and actions appreciated by horses. An overview of snort production in 127 horses spread over 16 riding schools was first conducted to highlight a potential site effect. Results show a great difference in snorts frequency between facilities which may be due to different riding techniques. In order to test this hypothesis, we then focused on 37 horse-rider dyads by scoring horses' postures (neck) and riders' positions (hand, reins) during, but also out of the context of snort production. Results show that snorts were particularly associated with phases when the rider technique, i.e. long and loose reins, allowed more comfort for the horse, especially while walking. Results were more mitigated for higher paces since the association of snorts with signs of comfort was less clear-cut. Snorts could therefore be useful tools for identifying better practices, especially at slower gaits. However, care has to be taken at higher paces.

Mutual interactions between cognition and welfare: The horse as an animal model.

Research in cognitive psychology has repeatedly shown how much cognition and emotions are mutually related to one another. Psychological disorders are associated with cognitive (attention, memory and judgment) biases and chronic pain may affect attention, learning or memory. Laboratory studies have provided useful insights about the processes involved but observations about spontaneous animal models, living in different stress/welfare conditions may help understand further how cognition and welfare are interrelated in the « real world ¼. Domestic horses constitute such a model as they live in a variety of conditions that impact differently their welfare state. In the present review, we try and provide an overview of the scientific literature on cognition and welfare of domestic horses and their interrelationship. We address how emotions and welfare may affect cognitive processes in horses and impact the way they perceive their environment (including work). We propose new methods for assessing the relationship between welfare and cognition and open up the discussion on the evolution of the brain and the part domestication may have played.

Influence of theatre hall layout on actors' and spectators' emotions.

"Audience effect" is the influence of an audience size or composition on the emotional state of a public speaker. One characteristic of the audience which has received little attention is the spatial position of observers. We tested the influence of three positions (frontal, bi-frontal, and quadri-frontal) on actors and spectators' emotions in real theatrical representations. Measurements consisted in self-report questionnaires and galvanic skin responses. The layout of the theatre hall influenced both cognitive and physiological components of emotions. Actors were more influenced than spectators and showed an overall accuracy in self-perception. The quadri-frontal audience received the highest scores in actors' feeling assessments and galvanic skin responses. In addition, we found a discrepancy between self-assessment of emotional states by spectators and how actors perceive them. Attention should thus be paid in the layout of performance places with obviously more attention from the public and better feelings for actors in more dispersed settings.

Attentional state and brain processes: state-dependent lateralization of EEG profiles in horses.

Lateralization of brain functions has been suggested to provide individuals with advantages, such as an increase of neural efficiency. The right hemisphere is likely to be specialized for processing attention for details and the left hemisphere for categorization of stimuli. Thus attentional processes actually may underlie lateralization. In the present study, we hypothesized that the attentional state of horses could be reflected in the lateralization of brain responses. We used i) a recently developed attention test to measure horses' visual attentional responses towards a standardized stimulus and ii) a recently developed portable EEG telemetric tool to measure brain responses. A particular emphasis was given to the types of waves (EEG power profile) and their side of production when horses were either attentive towards a visual stimulus or quiet standing. The results confirmed that a higher attentional state is associated with a higher proportion of gamma waves. There was moreover an interaction between the attentional state, the hemisphere and the EEG profile: attention towards the visual stimulus was associated with a significant increase of gamma wave proportion in the right hemisphere while "inattention" was associated with more alpha and beta waves in the left hemisphere. These first results are highly promising and contribute to the large debate on functional lateralization.

Spontaneous attention-capture by auditory distractors as predictor of distractibility: a study of domestic horses (Equus caballus).

Distractibility (i.e. individual distraction from his ongoing activity) is thoughts to affect daily life. The present study develops an easy way to assess inter-individual variations of distractibility of an animal model, the domestic horse. We developed the 'distractibility test' (DT), based on auditory stimuli, a major source of distraction in daily life. We hypothesized that the broadcast of unusual sounds would provide a reliable source of distraction and that the responses to these unusual sounds would yield a good estimation of a horse's level of distractibility. Validity of the DT was assessed by comparing the subjects' interest towards the sound in this test to their attentional state in experimental visual attention tasks and in a working task. Our results showed inter-individual differences in response to the stimuli, with consistency over time. The subjects' responses to this DT were negatively correlated to their attentional skills in separate experimental tests and in a working task. This is to our knowledge the first 'real-world' estimate of an animal's distractibility in its home environment that could potentially be adapted for humans.

A novel test for evaluating horses' spontaneous visual attention is predictive of attention in operant learning tasks.

Attention is described as the ability to process selectively one aspect of the environment over others. In this study, we characterized horses' spontaneous attention by designing a novel visual attention test (VAT) that is easy to apply in the animal's home environment. The test was repeated over three consecutive days and repeated again 6 months later in order to assess inter-individual variations and intra-individual stability. Different patterns of attention have been revealed: 'overall' attention when the horse merely gazed at the stimulus and 'fixed' attention characterized by fixity and orientation of at least the visual and auditory organs towards the stimulus. The individual attention characteristics remained consistent over time (after 6 months, Spearman correlation test, P < 0.05). The validity of this novel test as a predictor of individual attentional skills was assessed by comparing the results, for the same horses, with those obtained in both a 'classical' experimental attention test the 'five-choice serial reaction time task' (5-CSRTT) and a work situation (lunge working context). Our results revealed that (i) individual variations remained consistent across tests and (ii) the VAT attention measures were not only predictive of attentional skills but also of learning abilities. Differences appeared however between the first day of testing and the following test days: attention structure on the second day was predictive of learning abilities, attention performances in the 5-CSRRT and at work. The VAT appears as a promising easy-to-use tool to assess animals' attention characteristics and the impact of different factors of variation on attention.

Do horses with poor welfare show 'pessimistic' cognitive biases?

This field study tested the hypothesis that domestic horses living under putatively challenging-to-welfare conditions (for example involving social, spatial, feeding constraints) would present signs of poor welfare and co-occurring pessimistic judgement biases. Our subjects were 34 horses who had been housed for over 3 years in either restricted riding school situations (e.g. kept in single boxes, with limited roughage, ridden by inexperienced riders; N = 25) or under more naturalistic conditions (e.g. access to free-range, kept in stable social groups, leisure riding; N = 9). The horses' welfare was assessed by recording health-related, behavioural and postural indicators. Additionally, after learning a location task to discriminate a bucket containing either edible food ('positive' location) or unpalatable food ('negative' location), the horses were presented with a bucket located near the positive position, near the negative position and halfway between the positive and negative positions to assess their judgement biases. The riding school horses displayed the highest levels of behavioural and health-related problems and a pessimistic judgment bias, whereas the horses living under more naturalistic conditions displayed indications of good welfare and an optimistic bias. Moreover, pessimistic bias data strongly correlated with poor welfare data. This suggests that a lowered mood impacts a non-human species' perception of its environment and highlights cognitive biases as an appropriate tool to assess the impact of chronic living conditions on horse welfare.

Anesthesia and brain sensory processing: impact on neuronal responses in a female songbird.

Whether anesthesia impacts brain sensory processing is a highly debated and important issue. There is a general agreement that anesthesia tends to diminish neuronal activity, but its potential impact on neuronal "tuning" is still an open question. Here we show, based on electrophysiological recordings in the primary auditory area of a female songbird, that anesthesia induces neuronal responses towards biologically irrelevant sounds and prevents the seasonal neuronal tuning towards functionally relevant species-specific song elements.

Lower attention to daily environment: a novel cue for detecting chronic horses' back pain?

Chronic pain is thought to affect patients' cognitive functioning, including attention. Loss of attention is likely to have an impact on the execution of daily tasks, and, therefore, to have negative effects. However, relationships between chronic pain and cognitive deficits are still debated. Pre-clinical studies using laboratory animals prove useful to model pain-related cognitive impairment, but animal models had to predict effects in the real world. This study investigates attentional engagement of domestic horses by comparing observations in a home setting and evaluations of vertebral disorders. We found that lower attentional engagement and the level of back disorders were correlated. Two different evaluation techniques of the state of horses' spines gave similar results. We suggest that novel animal models would prove useful for identifying spontaneous behaviours indicative of chronic pain. We suggest that more ethological studies in human patients' home environments would help to improve our understanding of the processes involved. Finally, these results yield interesting indications for evaluating animal welfare, as attentional engagement could become a reliable indicator of chronic pain and thus a useful tool for identification of suffering individuals.

Investigating attentional processes in depressive-like domestic horses (Equus caballus).

Some captive/domestic animals respond to confinement by becoming inactive and unresponsive to external stimuli. Human inactivity is one of the behavioural markers of clinical depression, a mental disorder diagnosed by the co-occurrence of symptoms including deficit in selective attention. Some riding horses display 'withdrawn' states of inactivity and low responsiveness to stimuli that resemble the reduced engagement with their environment of some depressed patients. We hypothesized that 'withdrawn' horses experience a depressive-like state and evaluated their level of attention by confronting them with auditory stimuli. Five novel auditory stimuli were broadcasted to 27 horses, including 12 'withdrawn' horses, for 5 days. The horses' reactions and durations of attention were recorded. Non-withdrawn horses reacted more and their attention lasted longer than that of withdrawn horses on the first day, but their durations of attention decreased over days, but those of withdrawn horses remained stable. These results suggest that the withdrawn horses' selective attention is altered, adding to already evidenced common features between this horses' state and human depression.

Visual attention, an indicator of human-animal relationships? A study of domestic horses (Equus caballus).

As visual attention is an intrinsic part of social relationships, and because relationships are built on a succession of interactions, their establishment involves learning and attention. The emotional, rewarding or punishing, content can modulate selective attention. In horses, the use of positive/negative reinforcement during training determines short and long-term human-horse relationships. In a recent study in horses, where either food or withers' grooming were used as a reward, it appeared that only the food-rewarded horses learned the task and show better relationship with humans. In the present study, we hypothesized that this differential effect of grooming/food rewards on learning performances could be due to attentional processes. Monitoring, gazes and behaviors directed towards the trainer revealed that the use of a food reward (FR) as positive reinforcement increased horses' selective attention towards their trainer. Conversely, horses trained with grooming reward (GR) expressed more inattentive responses and did not show a decrease of "agitated" behavior. However, individual plotting of attention vs. rate of learning performances revealed a complex pattern. Thus, while all FR horses showed a "window" of attention related to faster learning performances, GR horses' pattern followed an almost normal curve where the extreme animals (i.e., highest and lowest attention) had the slowest learning performances. On the other hand, learning was influenced by attention: at the end of training, the more attentive horses had also better learning performances. This study, based on horses, contributes to the general debate on the place of attentional processes at the interface of emotion and cognition and opens new lines of thought about individual sensitivities (only individuals can tell what an appropriate reward is), attentional processes and learning.

Songs of two starling species: common traits versus adaptations to the social environment.

We analysed, for the first time, songs of the African Red-winged Starling Onychognathus morio and compared their general characteristics with those of the European Starling Sturnus vulgaris. Both species are gregarious during the non-breeding season, but European Starlings tend to nest in colonies, form unstable pair-bonds and are occasionally polygynous, whereas Red-winged Starlings form long-term pair-bonds and occupy exclusive nesting territories. Red-winged Starlings produced the same basic song categories as European Starlings: warbles and whistles. These two categories appeared to be involved in similar social interactions in the two species. However, several aspects of song behaviour differed between the two species: Red-winged Starlings, breeding in isolated nests, preferentially used whistles for long-distance communication and showed a simpler organization of warbling song. Whistles in the Red-winged Starling were mostly shared between birds and, in contrast to the European Starling, were not indicators of individual identity. Also in contrast to the European Starling, female song in Red-winged Starlings appeared very important throughout the breeding period. Our results suggest that some song characteristics in the two species are phylogenetically conserved whereas others are affected by the distinct social systems of the two species.

Do horses expect humans to solve their problems?

Domestic animals are highly capable of detecting human cues, while wild relatives tend to perform less well (e.g., responding to pointing gestures). It is suggested that domestication may have led to the development of such cognitive skills. Here, we hypothesized that because domestic animals are so attentive and dependant to humans' actions for resources, the counter effect may be a decline of self sufficiency, such as individual task solving. Here we show a negative correlation between the performance in a learning task (opening a chest) and the interest shown by horses toward humans, despite high motivation expressed by investigative behaviors directed at the chest. If human-directed attention reflects the development of particular skills in domestic animals, this is to our knowledge the first study highlighting a link between human-directed behaviors and impaired individual solving task skills (ability to solve a task by themselves) in horses.

Acoustic variability and social significance of calls in female Campbell's monkeys (Cercopithecus campbelli campbelli).

Although the vocal repertoire of nonhuman primates is strongly constrained by genetic, a growing number of studies evidence socially determined flexibility. According to Snowdon et al. [Social Influences on Vocal Development (University Press, Cambridge, 1997), pp. 234-248], calls with a higher social function (affiliative or agonistic) would be expected to show more flexibility than lesser social calls. Owren and Rendall [Evol. Anthropol., 10, 58-71 (2001)] nuanced this by defending a structure-function relationship. Calls with particular acoustic properties, which directly influence the listener's affect, would be less individually distinctive than calls involved in an affective conditioning process. These hypotheses were tested in Campbell's monkeys using telemetric recordings. This is the first detailed description of female Campbell's monkeys' vocal repertoire emphasizing a possible relationship between social function and flexibility level. The vocal repertoire displayed an "arborescent" organization (call type, subtype, and variants). The highest number of subtypes and the greatest acoustic variability, within and among individuals, were found in calls associated with the highest affiliative social value. However, calls associated with agonism were the most stereotyped, whereas less social alarm calls were intermediate. This only partially validate the hypothesis of Snowdon et al. In accordance with Owren and Rendall's hypotheses, the level of individual distinctiveness was minimum for noisy pulsed calls and maximum for calls involved in affiliative interactions.

Foraging opportunity: a crucial criterion for horse welfare?

This study aimed at determining the effect of the increase of foraging opportunities on the behaviour and welfare of breeding mares housed in individual boxes but allowed outside 6 h a day in a bare paddock. One hundred Arab breeding mares were divided into two groups of 50 according to the treatment and allowed outside in two bare paddocks at the same density (115 mare/ha) where water and shelter were provided. The treatment consisted in providing the opportunity to forage on hay. Twenty-minute animal focal samplings and scan samplings were used to determine the time budget of the mares during the period from 0900 to 1500 h and study their social behaviour. A total of 300 focal sampling (6000 min), 3300 individual scan sampling (6000 min) and 62 group observations (1240 min) corresponding to the 100 mares were recorded. Non-parametric tests were used to analyse data. Results showed that experimental mares spent more time feeding (65.12% ± 2.40% v. 29.75% ± 2.45%, P < 0.01) and less time in locomotion (11.70% ± 1.31% v. 23.56% ± 1.34%, P < 0.01), stand resting (11.76% ± 2.57% v. 27.52% ± 2.62%, P < 0.01) and alert standing (5.23% ± 1.2% v. 14.71% ± 1.23%, P < 0.01). There was more bonding among experimental mares than control ones (26 v. 14, P < 0.05). Experimental mares showed more positive social interactions (P < 0.01) and less aggression (P < 0.01). These results suggest that giving densely housed mares foraging opportunities improves their welfare.

Are 6-month-old foals sensitive to dam's influence?

A recent study has shown that gently handling dams in front of their few days old foals may strongly influence the development of human-foal relationships. In the present study, we test whether 6-month-old foals remain sensitive to their dams' influence. The study was performed on 16 foal-mare dyads, with half of the mares receiving positive contacts from the experimenter in presence of their 6-month-old foals (n = 8) whereas the other mares were not handled (n = 8). All foals were tested 15 and 30-35 days later under various conditions (reaction to a motionless human, approach test, saddle-pad tolerance test). We observe a positive effect of mare' handling on foals' reactions to humans but with a high interindividual variability, suggesting a higher effect of the foals' own behavioral characteristics at this age than at earlier stages.

Influence of various early human-foal interferences on subsequent human-foal relationship.

Whereas the way animals perceive human contact has been particularly examined in pet animals, a small amount of investigations has been done in domestic ungulates. It was nevertheless assumed that, as pet animals, non-aggressive forms of tactile contact were as well rewarding or positive for these species, even though the features of intraspecific relationships in pet animals and domestic ungulates may be to some extent different. We test here the hypothesis that horses may not consider physical handling by humans as a positive event. When comparing different early human-foal interactions, we found that early exposure to a motionless human enhanced slightly foals reactions to humans whereas forced stroking or handling in early life did not improve later human-foal relation. Foals that were assisted during their first suckling (e.g., brought to the dam's teat) even tended to avoid human approach at 2 weeks, and physical contact at 1 month of age. We argue that interspecies differences may exist in how tactile stimulation is perceived. It may be important for the establishment of a bond that a young animal is active in the process and able, through its behavioral responses, to help define what is positive for it. This way of investigation may have important general implications in how we consider the development of social relations, both within and between species.

New insights into the auditory processing of communicative signals in the HVC of awake songbirds.

In the present study, using a systematic recording method and a variety of stimuli, we determined the proportion of responsive sites and their response features in the vocal control nucleus HVC of awake-restrained starlings, a species with multiple song types. Responsive sites were classified into three groups, according to the number of stimuli to which they responded. Sites in the three groups showed responses to individual-specific songs, with sites in the group that showed responses to only one stimulus responding mostly to a bird's own song. In comparison, very few sites exhibited responses to universal species-specific songs and to artificial nonspecific sounds. By contrast, data obtained in the same birds under urethane anesthesia show that, although the total proportion of responsive sites was similar, numerous responses to a universal species-specific song and to an artificial nonspecific pure tone could be observed.

Functional organization of the forebrain auditory centres of the European starling: a study based on natural sounds.

The field L complex is thought to be the highest auditory centre and the input in the song vocal nuclei. Different anatomical and functional subdivisions have been described in field L. Auditory neurons of field L are well activated by natural sounds and especially by species-specific sounds. A complex sound coding appears to exist in field L. However, until now, the spatial organization of the different functional subdivisions has been described only using artificial sounds. Here, we investigated the spatial distribution of neuronal responses in field L to species-specific songs. Starlings seemed to be a very appropriate species for this investigation, both because of their complex vocal behaviour that implies different levels of categorization and their neuronal responses towards complex song elements. Multi-unit recordings were performed in wild starlings that were awake. The method of backward correlation was used to visualize the functional organization and we represented the neuronal responses as both activity maps and correlation maps. The use of natural sounds allowed us to define several functional sub-areas with different neuronal processing. These results show that field L is involved in a more complex task than simple frequency processing.