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.

Automatic acoustic identification of individuals in multiple species: improving identification across recording conditions.

Many animals emit vocal sounds which, independently from the sounds' function, contain some individually distinctive signature. Thus the automatic recognition of individuals by sound is a potentially powerful tool for zoology and ecology research and practical monitoring. Here, we present a general automatic identification method that can work across multiple animal species with various levels of complexity in their communication systems. We further introduce new analysis techniques based on dataset manipulations that can evaluate the robustness and generality of a classifier. By using these techniques, we confirmed the presence of experimental confounds in situations resembling those from past studies. We introduce data manipulations that can reduce the impact of these confounds, compatible with any classifier. We suggest that assessment of confounds should become a standard part of future studies to ensure they do not report over-optimistic results. We provide annotated recordings used for analyses along with this study and we call for dataset sharing to be a common practice to enhance the development of methods and comparisons of results.

Correction: The assessment of biases in the acoustic discrimination of individuals.

[This corrects the article DOI: 10.1371/journal.pone.0177206.].

Ontogeny of individual and litter identity signaling in grunts of piglets.

Many studies have shown that animal vocalizations can signal individual identity and group/family membership. However, much less is known about the ontogeny of identity information-when and how this individual/group distinctiveness in vocalizations arises and how it changes during the animal's life. Recent findings suggest that even species that were thought to have limited vocal plasticity could adjust their calls to sound more similar to each other within a group. It has already been shown that sows can acoustically distinguish their own offspring from alien piglets and that litters differ in their calls. Surprisingly, individual identity in piglet calls has not been reported yet. In this paper, this gap is filled, and it is shown that there is information about piglet identity. Information about litter identity is confirmed as well. Individual identity increased with age, but litter vocal identity did not increase with age. The results were robust as a similar pattern was apparent in two situations differing in arousal: isolation and back-test. This paper argues that, in piglets, increased individual discrimination results from the rapid growth of piglets, which is likely to be associated with growth and diversification of the vocal tract rather than from social effects and vocal plasticity.

The assessment of biases in the acoustic discrimination of individuals.

Animal vocalizations contain information about individual identity that could potentially be used for the monitoring of individuals. However, the performance of individual discrimination is subjected to many biases depending on factors such as the amount of identity information, or methods used. These factors need to be taken into account when comparing results of different studies or selecting the most cost-effective solution for a particular species. In this study, we evaluate several biases associated with the discrimination of individuals. On a large sample of little owl male individuals, we assess how discrimination performance changes with methods of call description, an increasing number of individuals, and number of calls per male. Also, we test whether the discrimination performance within the whole population can be reliably estimated from a subsample of individuals in a pre-screening study. Assessment of discrimination performance at the level of the individual and at the level of call led to different conclusions. Hence, studies interested in individual discrimination should optimize methods at the level of individuals. The description of calls by their frequency modulation leads to the best discrimination performance. In agreement with our expectations, discrimination performance decreased with population size. Increasing the number of calls per individual linearly increased the discrimination of individuals (but not the discrimination of calls), likely because it allows distinction between individuals with very similar calls. The available pre-screening index does not allow precise estimation of the population size that could be reliably monitored. Overall, projects applying acoustic monitoring at the individual level in population need to consider limitations regarding the population size that can be reliably monitored and fine-tune their methods according to their needs and limitations.

The traits of the great calls in the juvenile and adolescent gibbon males Nomascus gabriellae.

Knowledge about vocal ontogeny and vocal plasticity during ontogeny in primate species is central to understanding the evolution of human speech. Vocalizations in gibbons (Hominoidea) are very interesting and contain complex species- and sex-specific patterns. However, ontogeny of gibbon songs is little studied. Here, we document regular production and ontogenetic changes of female-specific "great call" in 4 immature (2 juvenile-c.a. 3 years old; and 2 adolescent-c.a. 5 years old) males of southern yellow-cheeked gibbon (N. gabriellae) over nine months. None of the males produced fully developed adult-like "great call" and little ontogenetic changes to "great calls" occurred. "Great calls" of sons were shorter, started higher and ended lower than those of their mothers. Regular production of twitter part of great call likely appears around 4th year as it was observed in adolescent but not in juvenile males.

Expression of Emotional Arousal in Two Different Piglet Call Types.

Humans as well as many animal species reveal their emotional state in their voice. Vocal features show strikingly similar correlation patterns with emotional states across mammalian species, suggesting that the vocal expression of emotion follows highly conserved signalling rules. To fully understand the principles of emotional signalling in mammals it is, however, necessary to also account for any inconsistencies in the way that they are acoustically encoded. Here we investigate whether the expression of emotions differs between call types produced by the same species. We compare the acoustic structure of two common piglet calls-the scream (a distress call) and the grunt (a contact call)-across three levels of arousal in a negative situation. We find that while the central frequency of calls increases with arousal in both call types, the amplitude and tonal quality (harmonic-to-noise ratio) show contrasting patterns: as arousal increased, the intensity also increased in screams, but not in grunts, while the harmonicity increased in screams but decreased in grunts. Our results suggest that the expression of arousal depends on the function and acoustic specificity of the call type. The fact that more vocal features varied with arousal in scream calls than in grunts is consistent with the idea that distress calls have evolved to convey information about emotional arousal.

Humans (Homo sapiens) judge the emotional content of piglet (Sus scrofa domestica) calls based on simple acoustic parameters, not personality, empathy, nor attitude toward animals.

The vocal expression of emotion is likely driven by shared physiological principles among species. However, which acoustic features promote decoding of emotional state and how the decoding is affected by their listener's psychology remain poorly understood. Here we tested how acoustic features of piglet vocalizations interact with psychological profiles of human listeners to affect judgments of emotional content of heterospecific vocalizations. We played back 48 piglet call sequences recorded in four different contexts (castration, isolation, reunion, nursing) to 60 listeners. Listeners judged the emotional intensity and valence of the recordings and were further asked to attribute a context of emission from four proposed contexts. Furthermore, listeners completed a series of questionnaires assessing their personality (NEO-FFI personality inventory), empathy [Interpersonal Reactivity Index (IRI)] and attitudes to animals (Animal Attitudes Scale). None of the listeners' psychological traits affected the judgments. On the contrary, acoustic properties of recordings had a substantial effect on ratings. Recordings were rated as more intense with increasing pitch (mean fundamental frequency) and increasing proportion of vocalized sound within each stimulus recording and more negative with increasing pitch and increasing duration of the calls within the recording. More complex acoustic properties (jitter, harmonic-to-noise ratio, and presence of subharmonics) did not seem to affect the judgments. The probability of correct context recognition correlated positively with the assessed emotion intensity for castration and reunion calls, and negatively for nursing calls. In conclusion, listeners judged emotions from pig calls using simple acoustic properties and the perceived emotional intensity might guide the identification of the context.

Being angry, singing fast? Signalling of aggressive motivation by syllable rate in a songbird with slow song.

Syllable rate has been shown to play a role in male-male aggressive interactions and has been proposed to serve as a male quality indicator in several bird species. In those with fast syllable rates, males often increase rates when singing in aggressive context, and respond differently to test stimuli of varying rates. We asked whether the syllable rate fulfils a similar signalling function in the chiffchaff (Phylloscopus collybita), a songbird species with a slow syllable rate. We confronted 36 chiffchaff males with one of three playback types differing in syllable rate: control (non-manipulated rate), fast, or slow (artificially increased and decreased syllable rate, respectively). We recorded tested males' songs and behaviour before and during the experiment. Our results indicate that syllable rate might be an aggressive signal in chiffchaff. Males that physically attacked the loudspeaker during experiments sang faster songs spontaneously, and those that continued singing during the playback responded to fast and non-manipulated stimuli with substantial increase of syllable rate. Indirect evidence further suggests that syllable rate in chiffchaff is unlikely constrained by respiratory demands; thus, we propose that syllable rate in this species functions as a conventional signal of male aggressiveness rather than an index of quality.

Encoding of situations in the vocal repertoire of piglets (Sus scrofa): a comparison of discrete and graded classifications.

Two important questions in bioacoustics are whether vocal repertoires of animals are graded or discrete and how the vocal expressions are linked to the context of emission. Here we address these questions in an ungulate species. The vocal repertoire of young domestic pigs, Sus scrofa, was quantitatively described based on 1513 calls recorded in 11 situations. We described the acoustic quality of calls with 8 acoustic parameters. Based on these parameters, the k-means clustering method showed a possibility to distinguish either two or five clusters although the call types are rather blurred than strictly discrete. The division of the vocal repertoire of piglets into two call types has previously been used in many experimental studies into pig acoustic communication and the five call types correspond well to previously published partial repertoires in specific situations. Clear links exist between the type of situation, its putative valence, and the vocal expression in that situation. These links can be described adequately both with a set of quantitative acoustic variables and through categorisation into call types. The information about the situation of emission of the calls is encoded through five call types almost as accurately as through the full quantitative description.