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Oestrus detection remains a problem in the dairy cattle industry. Therefore, automatic detection systems have been developed to detect specific behavioural changes at oestrus. Vocal behaviour has not been considered in such automatic oestrus detection systems in cattle, though the vocalisation rate is known to increase during oestrus. The main challenge in using vocalisation to detect oestrus is correctly identifying the calling individual when animals are moving freely in large groups, as oestrus needs to be detected at an individual level. Therefore, we aimed to automate vocalisation recording and caller identification in group-housed dairy cows. This paper first presents the details of such a system and then presents the results of a pilot study validating its functionality, in which the automatic detection of calls from individual heifers was compared to video-based assessment of these calls by a trained human observer, a technique that has, until now, been considered the ‘gold standard’. We developed a collar-based cattle call monitor (CCM) with structure-borne and airborne sound microphones and a recording unit and developed a postprocessing algorithm to identify the caller by matching the information from both microphones. Five group-housed heifers, each in the perioestrus or oestrus period, were equipped with a CCM prototype for 5 days. The recorded audio data were subsequently analysed and compared with audiovisual recordings. Overall, 1404 vocalisations from the focus heifers and 721 vocalisations from group mates were obtained. Vocalisations during collar changes or malfunctions of the CCM were omitted from the evaluation. The results showed that the CCM had a sensitivity of 87% and a specificity of 94%. The negative and positive predictive values were 80% and 96%, respectively. These results show that the detection of individual vocalisations and the correct identification of callers are possible, even in freely moving group-housed cattle. The results are promising for the future use of vocalisation in automatic oestrus detection systems.

Good management and improved standards of animal welfare are discussed as important ways of reducing the risk of infection in farm animals without medication. Increasing evidence from both humans and animals suggests that environments that promote wellbeing over stress and positive over negative emotions can reduce susceptibility to disease and/or lead to milder symptoms. We point out, however, that the relationship between welfare, immunity, and disease is highly complex and we caution against claiming more than the current evidence shows. The accumulating but sometimes equivocal evidence of close links between the brain, the gut microbiome, immunity, and welfare are discussed in the context of the known links between mental and physical health in humans. This evidence not only provides empirical support for the importance of good welfare as preventative medicine in animals but also indicates a variety of mechanisms by which good welfare can directly influence disease resistance. Finally, we outline what still needs to be done to explore the potential preventative effects of good welfare., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Düpjan and Dawkins.)

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., (© 2022. The Author(s).)

Both humans and nonhuman animals need to show self-control and wait for a larger or better reward instead of a smaller or less preferred but instant reward on an everyday basis. We investigated whether this ability undergoes ontogenetic development in domestic pigs (similar to what is known in human infants) by testing if and for how long nine- and 16-week-old pigs wait for a larger amount of their preferred reward. In a delay-of-gratification task, animals first learned that a small reward was hidden under a white cup and a large reward under a black cup, and then the delay to deliver the large reward was gradually increased. The results show that older pigs could wait longer for a larger reward than younger pigs (10.6 ± 1.3 s vs. 5.2 ± 1.5 s), thereby confirming our hypothesis of ontogenetic development of self-control in pigs. This self-control is likely to be regulated by the behavioural inhibition system and associated systems. Self-control or, more specifically the lack of it may be involved in the development of abnormal behaviours, not only in humans but also in animals. Therefore, research on self-control in decision-making might provide a new perspective on abnormal behaviours in captive animals., (© 2021. The Author(s).)

In humans, speech perception is lateralized, with the left hemisphere of the brain dominant in processing the communicative content and the right hemisphere dominant in processing the emotional content. However, still little is known about such a division of tasks in other species. We therefore investigated lateralized processing of communicative and emotionally relevant calls in a social mammal, the pig ( Sus scrofa ). Based on the contralateral connection between ears and hemispheres, we compared the behavioural and cardiac responses of 36 young male pigs during binaural and monaural (left or right) playback to the same sounds. The playback stimuli were calls of social isolation and physical restraint, whose communicative and emotional relevance, respectively, were validated prior to the test by acoustic analyses and during binaural playbacks. There were indications of lateralized processing mainly in the initial detection (left head-turn bias, indicating right hemispheric dominance) of the more emotionally relevant restraint calls. Conversely, there were indications of lateralized processing only in the appraisal (increased attention during playback to the right ear) of the more communicative relevant isolation calls. This implies differential involvement of the hemispheres in the auditory processing of vocalizations in pigs and thereby hints at similarities in the auditory processing of vocal communication in non-human animals and speech in humans. Therefore, these findings provide interesting new insight in the evolution of human language and auditory lateralization., (Copyright © 2020 Leliveld, Düpjan, Tuchscherer and Puppe.)

Oestrus detection remains a problem in the dairy cattle industry. Therefore, automatic detection systems have been developed to detect specific behavioural changes at oestrus. Vocal behaviour has not been considered in such automatic oestrus detection systems in cattle, though the vocalisation rate is known to increase during oestrus. The main challenge in using vocalisation to detect oestrus is correctly identifying the calling individual when animals are moving freely in large groups, as oestrus needs to be detected at an individual level. Therefore, we aimed to automate vocalisation recording and caller identification in group-housed dairy cows. This paper first presents the details of such a system and then presents the results of a pilot study validating its functionality, in which the automatic detection of calls from individual heifers was compared to video-based assessment of these calls by a trained human observer, a technique that has, until now, been considered the 'gold standard'. We developed a collar-based cattle call monitor (CCM) with structure-borne and airborne sound microphones and a recording unit and developed a postprocessing algorithm to identify the caller by matching the information from both microphones. Five group-housed heifers, each in the perioestrus or oestrus period, were equipped with a CCM prototype for 5 days. The recorded audio data were subsequently analysed and compared with audiovisual recordings. Overall, 1404 vocalisations from the focus heifers and 721 vocalisations from group mates were obtained. Vocalisations during collar changes or malfunctions of the CCM were omitted from the evaluation. The results showed that the CCM had a sensitivity of 87% and a specificity of 94%. The negative and positive predictive values were 80% and 96%, respectively. These results show that the detection of individual vocalisations and the correct identification of callers are possible, even in freely moving group-housed cattle. The results are promising for the future use of vocalisation in automatic oestrus detection systems.

Animal individuality is challenging to explain because individual differences are regulated by multiple selective forces that lead to unique combinations of characteristics. For instance, the study of personality, a core aspect of individuality, may benefit from integrating other factors underlying individual differences, such as lateralized cerebral processing. Indeed, the approach-withdrawal hypothesis (the left hemisphere controls approach behavior, the right hemisphere controls withdrawal behavior), may account for differences in boldness or exploration between left and right hemispheric dominant individuals. To analyze the relationships between personality and laterality we tested 80 male piglets with established laterality patterns for 2 motor functions (tail curling direction and the side of the snout used for manipulation) and a combined classification integrating both motor functions using cluster analysis. We analyzed basal salivary testosterone and cortisol along with their behavior in standardized tests as pre-established indicators of different personality traits (Boldness, Exploration, Activity, Sociability, and Coping). We found that the direction of the single motor biases showed significant associations with few personality traits. However, the combined laterality classification showed more, and more robust, significant associations with different personality traits compared with the single motor biases. These results supported the approach-withdrawal hypothesis because right-biased pigs were bolder and more explorative in a context of novelty. Additionally, right-biased pigs were more sociable than left-biased pigs. Therefore, the present study indicates that personality is indeed related to lateralized cerebral processing and provides insight into the multifactorial nature of individuality., (© The Author(s) (2018). Published by Oxford University Press.)

Motor lateralization is hypothesized to depend on the complexity of the motor function, but it might at the same time reflect hemispheric dominance within an individual across motor functions. We investigated possible motor lateralization patterns in four motor functions of different complexity (snout use in a manipulative task, foot use in two-stepping tasks and tail curling) in the domestic pig, a tetrapod species relevant as farm animal but also as a model in human neuroscience. A significant majority of our sample showed individual biases for manipulation with their snout and for curling their tail. Interestingly, the tail curling was lateralized towards the right at the population level and showed stronger lateralization patterns than the snout. Using a cluster analysis with combined tail and snout laterality, we identified groups of individuals with different lateralization patterns across motor functions that potentially reflect the individuals' hemispheric dominance. To conclude, our results suggest that pigs show lateralization patterns that depend on the motor function and on the individual. Such individual lateralization patterns might have broader implications for animal personality and welfare. Our study lays the methodological groundwork for future research on laterality in pigs.

The reliable detection of estrus is an important scientific and practical challenge in dairy cattle farming. Female vocalization may indicate reproductive status, and preliminary evidence suggests that this information can be used to detect estrus in dairy cattle. The aim of this study was to associate the changes in the vocalization rate of dairy heifers with behavioral estrus indicators as well as test the influence of the type of estrus (natural estrus vs. superovulation-induced estrus). We analyzed 6 predefined estrus-related behavior patterns (standing to be mounted, head-side mounting, active mounting, chin resting, being mounted while not standing, and active sniffing in the anogenital region) and vocalization rates in the peri-estrus period (day of estrus ± 1 d) of 12 German Holstein heifers using audio-visual recordings. Each heifer was observed under natural estrus and a consecutive superovulation induced by FSH and cloprostenol. Estrus was determined by behavioral patterns and confirmed by clinical examination (vaginoscopy and ultrasound imaging of the ovaries) as well as by the concentration of peripheral progesterone. Estrus behavior and vocalization rates were analyzed in 3-h intervals (an average of 19 intervals for each heifer), and an estrus score was calculated based on the 6 behaviors. The interval with the highest estrus score (I0) was considered the estrus climax. We demonstrated similar time courses for the estrus score and vocalization rate independent of estrus type. However, in natural estrus, the maximum vocalization rate (±SE) occurred in the interval before estrus climax (I-1; 42.58 ± 21.89) and was significantly higher than that in any other interval except estrus climax (I0; 27.58 ± 9.76). During natural estrus, the vocalization rate was significantly higher within the interval before estrus climax (I-1; 42.58 ± 21.89 vs. 11.58 ± 5.51) than under superovulation. The results underscore the potential use of vocalization rate as a suitable indicator of estrus climax in automated estrus detection devices. Further studies and technical development are required to record and process individual vocalization rates., (The Authors. Published by FASS Inc. and Elsevier Inc. on behalf of the American Dairy Science Association®. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/3.0/).)

Vocalizations have long been recognized to encode information about an individual's emotional state and, as such, have contributed to the study of emotions in animals. However, the potential of vocalizations to also encode information about an individual's emotional reactivity has received much less attention. In this study, we aimed to test whether the vocalizations of domestic pigs contain correlates of emotional reactivity that are consistent between different contexts. We recorded vocalizations of 120 young female pigs in an experimental arena in two consecutive recording contexts, social isolation and an encounter with a familiar human. Simultaneously, we measured their heart rate and behaviour to determine their emotional reactivity in the same context (within-context). In addition, we aimed to determine the subjects' emotional reactivity in other contexts (across-context) by measuring their behaviour in four common tests of emotional reactivity, the human approach test, the open door test, the open field test and the novel object test. Using a cluster analysis, we identified four different call types. Significant inter-context correlations were found for all call types, suggesting that pig vocalizations are consistent within an individual across contexts. The call rate and the proportions of the individual call types were found to correlate significantly with indices of emotional reactivity both within and across contexts. Thereby, we found more significant correlations to indices of emotional reactivity within context (behavioural and physiological response during recording) compared to across context (behavioural response in the four emotional reactivity tests). The consistency of the vocal correlates to emotional reactivity between the different contexts depended on the call type. While we found moderate evidence that the high grunt is indicative of more active, more explorative and less fearful individuals both within as well as across contexts, the other call types provided less consistent results. Thus, it seems that some call types are better suited to provide information on a caller's emotional reactivity than others, and further research is needed to clarify the underlying influential factors., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Cognitive and affective processes are highly interrelated. This has implications for neuropsychiatric disorders such as major depressive disorder in humans but also for the welfare of non-human animals. The brain serotonergic system might play a key role in mediating the relationship between cognitive functions and affective regulation. The aim of our study was to examine the influence of serotonin depletion on the affective state and cognitive processing in pigs, an important farm animal species but also a potential model species for biomedical research in humans. For this purpose, we modified a serotonin depletion model using para-chlorophenylalanine (pCPA) to decrease serotonin levels in brain areas involved in cognitive and affective processing (part 1). The consequences of serotonin depletion were then measured in two behavioral tests (part 2): the spatial judgement task (SJT), providing information about the effects of the affective state on cognitive processing, and the open field/novel object (OFNO) test, which measures behavioral reactions to novelty that are assumed to reflect affective state. In part 1, 40 pigs were treated with either pCPA or saline for six consecutive days. Serotonin levels were assessed in seven different brain regions 4, 5, 6, 11 and 13days after the first injection. Serotonin was significantly depleted in all analyzed brain regions up to 13days after the first application. In part 2, the pCPA model was applied to 48 animals in behavioral testing. Behavioral tests, the OFNO test and the SJT, were conducted both before and after pCPA/saline injections. While results from the OFNO tests were inconclusive, an effect of treatment as well as an effect of the phase (before and after treatment) was observed in the SJT. Animals treated with pCPA showed more pessimistic-like behavior, suggesting a more negative affective state due to serotonin depletion. Thus, our results confirm that the serotonergic system is a key player in cognitive-emotional processing. Hence, the serotonin depletion model and the spatial judgement task can increase our understanding of the basic mechanisms underlying both human neuropsychiatric disorders and animal welfare., (Copyright © 2017 Elsevier Inc. All rights reserved.)

In the study of animal emotions, emotional valence has been found to be difficult to measure. Many studies of farm animals' emotions have therefore focussed on the identification of indicators of strong, mainly negative, emotions. However, subtle variations in emotional valence, such as those caused by rather moderate differences in husbandry conditions, may also affect animals' mood and welfare when such variations occur consistently. In this study, we investigated whether repeated moderate aversive or rewarding events could lead to measurable differences in emotional valence in young, weaned pigs. We conditioned 105 female pigs in a test arena to either a repeated startling procedure (sudden noises or appearances of objects) or a repeated rewarding procedure (applesauce, toy and straw) over 11 sessions. Control pigs were also regularly exposed to the same test arena but without conditioning. Before and after conditioning, we measured heart rate and its variability as well as the behavioural reactions of the subjects in the test arena, with a special focus on detailed acoustic analyses of their vocalisations. The behavioural and heart rate measures were analysed as changes compared to the baseline values before conditioning. A limited number of the putative indicators of emotional valence were affected by the conditioning. We found that the negatively conditioned pigs showed changes that were significantly different from those in control pigs, namely a decrease in locomotion and an increase in standing. The positively conditioned pigs, however, showed a stronger increase in heart rate and a smaller decrease in SDNN (a heart rate variability parameter indicating changes in autonomic regulation) compared to the controls. Compared to the negatively conditioned pigs, the positively conditioned pigs produced fewer vocalisations overall as well as fewer low-frequency grunts but more high-frequency grunts. The low-frequency grunts of the negatively conditioned pigs also showed lower frequency parameters (bandwidth, maximum frequency, 25% and 50% quartiles) compared to those of the positively conditioned pigs. In any of the statistically significant results, the conditioning accounted for 1.5-11.9% of variability in the outcome variable. Hence, we conclude that repeated moderate aversive and rewarding events have weak but measurable effects on some aspects of behaviour and physiology in young pigs, possibly indicating changes in emotional valence, which could ultimately affect their welfare. The combination of ethophysiological indicators, i.e., the concurrent examination of heart rate measures, behavioural responses and especially vocalisation patterns, as used in the current study, might be a useful way of examining subtle effects on emotional valence in further studies., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Abnormal behaviour is a potential indicator of pain, suffering and injury in captive animals. Especially stereotypies, i. e. repetitive invariant behavioural patterns without obvious function or goal, can be observed as a consequence of inadequate housing conditions. Hence, they are often considered indicators of impaired welfare. In context of the ongoing scientific debate on captive animal welfare, the number of publications on stereotypies has increased, most notably in veterinary and farm animal research. Based on biological principles and definitions, we present several examples of stereotypic behaviour in (mainly) farm animals, and discuss approaches of preventing or reducing them. The occurrence of abnormal behaviour is often, but not necessarily, associated with the fact that modern housing and management precludes various evolutionary emerged highly motivated behaviours, or poses challenges the animals are unable to cope with adequately. Numerous studies show that stereotypies can be indicative of (current or past) suffering and impaired welfare. They can be avoided or at least reduced by increasing the biological relevance of the housing environments through environmental enrichment which stimulates species-specific behaviour.

In domestic pigs, vocalisation can be an indicator of distress and negative emotional states. It might play a role in the transfer of emotion between individuals ('emotional contagion' or 'empathy'), which could result in impaired animal welfare on a group level based on the distress in an individual member of the group. The aim of this study was to characterise the responses of pigs to conspecific distress calls. We performed a playback experiment in an open arena with 24 juvenile German Landrace pigs, during which individual subjects were exposed to both conspecific distress calls and an artificial sine tone (control) on consecutive experimental days. Both behavioural (locomotion, vocalisation, elimination and distance to the speakers) and physiological responses (heart rate and heart rate variability) were measured for 2 min before, during and after the playback of the stimuli (distress calls/control). Subjects showed decreased locomotion and vocalisation rates during both stimuli, suggesting that the animals responded to both stimuli. Heart rates decreased at the onset of both stimuli due to an activation of the parasympathetic system, indicating an orientation response to sudden stimuli. However, heart rates decreased after the end of the distress calls but not after control stimuli, illustrating that conspecific calls and other sounds are evaluated differently. We conclude that pigs exposed to isolation are attentive to conspecific distress vocalisation and hence the information about threat possibly conveyed in it, but they do not share the distress of the caller. Therefore, we could not find direct effects of distress calls of unfamiliar conspecifics on the welfare of isolated juvenile pigs. However, the state of heightened attention elicited by conspecific distress calls may affect a pig's subsequent evaluation of its environment., (Copyright © 2011 Elsevier Inc. All rights reserved.)

In a pilot study we have injected the amygdala of five female pigs (age 8 weeks) with acetylcholine (ACh, 5.5muM/20mul) and recorded short latency utterances. The evoked vocalizations displayed the characteristics of natural screams in sonagram appearance and hearing impression. Quantitative analyses, too, revealed the similarity of the ACh-evoked vocalizations with natural screaming. The results demonstrate for the first time that aversive vocalizations can be triggered by cholinergic amygdala stimulation.