Cross-species discrimination of vocal expression of emotional valence by Equidae and Suidae.

BACKGROUND Discrimination and perception of emotion expression regulate interactions between conspecifics and can lead to emotional contagion (state matching between producer and receiver) or to more complex forms of empathy (e.g., sympathetic concern). Empathy processes are enhanced by familiarity and physical similarity between partners. Since heterospecifics can also be familiar with each other to some extent, discrimination/perception of emotions and, as a result, emotional contagion could also occur between species. RESULTS Here, we investigated if four species belonging to two ungulate Families, Equidae (domestic and Przewalski's horses) and Suidae (pigs and wild boars), can discriminate between vocalizations of opposite emotional valence (positive or negative), produced not only by conspecifics, but also closely related heterospecifics and humans. To this aim, we played back to individuals of these four species, which were all habituated to humans, vocalizations from a unique set of recordings for which the valence associated with vocal production was known. We found that domestic and Przewalski's horses, as well as pigs, but not wild boars, reacted more strongly when the first vocalization played was negative compared to positive, regardless of the species broadcasted. CONCLUSIONS Domestic horses, Przewalski's horses and pigs thus seem to discriminate between positive and negative vocalizations produced not only by conspecifics, but also by heterospecifics, including humans. In addition, we found an absence of difference between the strength of reaction of the four species to the calls of conspecifics and closely related heterospecifics, which could be related to similarities in the general structure of their vocalization. Overall, our results suggest that phylogeny and domestication have played a role in cross-species discrimination/perception of emotions

Novel ideas to further expand the applicability of rhythm analysis

The temporal structure of animals’ acoustic signals can inform about context, urgency, species, individual identity, or geographical origin. We present three independent ideas to further expand the applicability of rhythm analysis for isochronous, that is, metronome-like, rhythms. A description of a rhythm or beat needs to include a description of its goodness of fit, meaning how well the rhythm describes a sequence. Existing goodness-of-fit values are not comparable between methods and datasets. Furthermore, they are strongly correlated with certain parameters of the described sequence, for example, the number of elements in the sequence. We introduce a new universal goodness-of-fit value, ugof, comparable across methods and datasets, which illustrates how well a certain beat frequency in Hz describes the temporal structure of a sequence of elements. We then describe two additional approaches to adapt already existing methods to analyze the rhythm of acoustic sequences of animals. The new additions, a slightly modified way to use the already established Fourier analysis and concrete examples on how to use the visualization with recurrence plots, enable the analysis of more variable data, while giving more details than previously proposed measures. New methods are tested on 6 datasets including the very complex flight songs of male skylarks. The ugof is the first goodness-of-fit value capable of giving the information per element, instead of only per sequence. Advantages and possible interpretations of the new approaches are discussed. The new methods enable the analysis of more variable and complex communication signals. They give indications on which levels and structures to analyze and enable to track changes and differences in individuals or populations, for instance, during ontogeny or across regions. Especially, the ugof is not restricted to the analysis of acoustic signals but could for example also be applied on heartbeat measurements. Taken together, the ugof and proposed method additions greatly broaden the scope of rhythm analysis methods

Goats distinguish between positive and negative emotion-linked vocalisations

Background Evidence from humans suggests that the expression of emotions can regulate social interactions and promote coordination within a group. Despite its evolutionary importance, social communication of emotions in non-human animals is still not well understood. Here, we combine behavioural and physiological measures, to determine if animals can distinguish between vocalisations linked to different emotional valences (positive and negative). Using a playback paradigm, goats were habituated to listen to a conspecific call associated with positive or negative valence (habituation phase) and were subsequently exposed to a variant of the same call type (contact call) associated with the opposite valence (dishabituation phase), followed by a final call randomly selected from the habituation phase as control (rehabituation phase). The effects of the calls on the occurrence of looking and cardiac responses in these phases were recorded and compared. Results We found that when the valence of the call variant changed, goats were more likely to look at the source of the sound, indicating that they could distinguish calls based on their valence. Heart rate was not affected by the valence of the calls played, whereas heart-rate variability tended to be higher in the habituation and rehabituation phases, when positive calls were played compared to negative ones. Together, the behavioural and physiological measures provide evidence suggesting, first, that goats are able to distinguish call variants based on their valence, and second, that goat behaviour and cardiac responses are affected by call valence. Conclusion This study indicates that auditory modalities are a potent means to communicate emotions in non-human animals. These findings can contribute to our understanding of the evolution of emotion perception in non-human animals.ISSN:1742-999

Behaviour of Horses in a Judgment Bias Test Associated with Positive or Negative Reinforcement

Moods can influence our judgment of ambiguous stimuli as positive or negative. Measuring judgment bias in animals is a promising method to objectively assess their emotional states. Our study aimed to develop a cognitive bias test in horses, in order to assess the effect of training using positive reinforcement (PR) or negative reinforcement (NR) on their emotional states. We trained 12 mares to discriminate between a rewarded and a non-rewarded location situated on each side of a paddock. The mares were then trained during five days to perform several exercises using PR (n = 6) for one group, and NR (n = 6) for the other (treatment). Finally, we compared the responses of the two groups to three ambiguous locations situated between the rewarded and non-rewarded locations (judgment bias test). During the training exercises, according to our predictions, behavioural measures suggested that NR mares experienced more negative emotions than PR mares. Surprisingly, the results of the judgment bias test suggest that NR mares were in a more optimistic mood compared to PR mares, despite previously experiencing more negative emotions during the treatment. NR mares could have been more motivated to obtain a food reward than PR mares, which had been rewarded throughout the treatment phase. Alternatively, NR mares could have developed optimistic bias triggered by release from the negative state experienced during treatment. This first attempt to test judgment bias in horses suggests that this is a promising method to measure horse mood. Knowledge about the effect of training methods on the mental health of domesticated animals can add a new dimension to animal welfare, in order to promote better ways to work with animals

Perceptual lateralization of vocal stimuli in goats

Functional asymmetries, for example, the preferential involvement of 1 brain hemisphere to process stimuli, may increase brain efficiency and the capacity to carry out tasks simultaneously. We investigated which hemisphere was primarily involved in processing acoustic stimuli in goats using a head-orienting paradigm. Three playbacks using goat vocalizations recorded in different contexts: food anticipation (positive), isolation (negative), food frustration (negative), as well as 1 playback involving dog barks (negative) were presented on the left and right sides of the test subjects simultaneously. The head-orienting response (left or right) and latency to resume feeding were recorded. The direction of the head-orienting response did not differ between the various playbacks. However, when the head-orienting response was tested against chance level, goats showed a right bias regardless of the stimuli presented. Goats responded more to dog barks than to food frustration calls, whereas responses to food anticipation and isolation calls were intermediate. In addition, the latency to resume feeding, an indicator of fear reaction, was not affected by the kind of vocalization presented. These results provide evidence for asymmetries in goat vocal perception of emotional-linked conspecific and heterospecific calls. They also suggest involvement of the left brain hemisphere for processing acoustic stimuli, which might have been perceived as familiar and non-threatening

Fallow bucks attend to vocal cues of motivation and fatigue

Breeding vocalizations can provide up-to-date information about callers. During the rut, fallow buck calling rates vary in response to their nearest neighbors, and vocal fatigue is linked to body condition loss. We found that fallow bucks perceive higher calling rates as a greater threat and that bucks were sensitive to declines in the quality of the calls of other males. Over the breeding season, fallow bucks gain information from changing calls to continually assess the condition and motivation of conspecific

Elevated sensitivity to tactile stimuli in stereotypic horses

Although stereotypic behaviours are a common problem in captive animals, why certain individuals are more prone to develop them remains elusive. In horses, individuals show considerable differences in how they perceive and react to external events, suggesting that this may partially account for the emergence of stereotypies in this species. In this study, we focussed on crib-biting, the most common stereotypy displayed by horses. We compared how established crib-biters (“CB”=19) and normal controls (“C”=18) differed in response to a standard ‘personality’ assessment test battery, i.e. reactivity to humans, tactile sensitivity, social reactivity, locomotor activity, and curiosity versus fearfulness (both in novel and suddenness situations). Our analyses showed that crib-biters only differed from control horses in their tactile sensitivity, suggesting an elevated sensitivity to tactile stimuli. We suggest that this higher tactile sensitivity could be due to altered dopamine or endogenous opioid physiology, resulting from chronic stress exposition. We discuss these findings in relation to the hypothesis that there may be a genetic predisposition for stereotypic behaviour in horses, and in relation to current animal husbandry and management practises.Publisher PDFPeer reviewe

Classification of pig calls produced from birth to slaughter according to their emotional valence and context of production

Vocal expression of emotions has been observed across species and could provide a non-invasive and reliable means to assess animal emotions. We investigated if pig vocal indicators of emotions revealed in previous studies are valid across call types and contexts, and could potentially be used to develop an automated emotion monitoring tool. We performed an analysis of an extensive and unique dataset of low (LF) and high frequency (HF) calls emitted by pigs across numerous commercial contexts from birth to slaughter (7414 calls from 411 pigs). Our results revealed that the valence attributed to the contexts of production (positive versus negative) affected all investigated parameters in both LF and HF. Similarly, the context category affected all parameters. We then tested two different automated methods for call classification; a neural network revealed much higher classification accuracy compared to a permuted discriminant function analysis (pDFA), both for the valence (neural network: 91.5%; pDFA analysis weighted average across LF and HF (cross-classified): 61.7% with a chance level at 50.5%) and context (neural network: 81.5%; pDFA analysis weighted average across LF and HF (cross-classified): 19.4% with a chance level at 14.3%). These results suggest that an automated recognition system can be developed to monitor pig welfare on-farm.publishedVersio