Thierry Aubin

Seewave: a free modular tool for sound analysis and synthesis.

ABSTRACT We review Seewave, new software for analysing and synthesizing sounds. Seewave is free and works on a wide variety of operating systems as an extension of the R operating environment. Its current 67 functions allow the user to achieve time, amplitude and frequency analyses, to estimate quantitative differences between sounds, and to generate new sounds for playback experiments. Thanks to its implementation in the R environment, Seewave is fully modular. All functions can be combined for complex data acquisition and graphical output, they can be part of important scripts for batch processing and they can be modified ad libitum. New functions can also be written, making Seewave a truly open-source tool.

Finding a parent in a king penguin colony: the acoustic system of individual recognition

To be fed, a king penguin, Aptenodytes patagonicus, chick must identify the call of its parents, in the continuous background noise of the colony. To study this recognition process, we played back to the chicks parental calls with acoustic parameters modified in the temporal and frequency domains. The parental call is composed of syllables (complex sounds with harmonic series) separated by pronounced amplitude declines. Our experiments with modified signals indicate that the chicks frequency analysis of the call is not tuned towards precise peak energy values, the signal being recognized even when the carrier frequency was shifted 100 Hz down or 75 Hz up. To recognize the adult, chicks used frequency rather than amplitude modulation, in particular the frequency modulation shape of the syllable. This structure is repeated through the different syllables of the call giving a distinct vocal signature. Our experiments also show that the receiver needs to perceive only a small part of the signal: the first half of the syllable (0.23 s) and the first three harmonics were sufficient to elicit recognition. The small amount of information necessary to understand the message, the high redundancy in the time and frequency domains and the almost infinite possibilities of coding provided by the frequency modulation signature permit the chick to recognize the adult, without the help of a nest site. For these reasons, the code used in the call of the king penguin can be regarded as a functional code, increasing the possibility of individual recognition in an acoustically constraining environment. Copyright 1999 The Association for the Study of Animal Behaviour.

How to vocally identify kin in a crowd: The penguin model

Summary On penguins, individual recognition is observed between mates and between parents and chick(s). During the past five years, their particular strategies of coding—decoding have been tested by playing back modified display calls to six species, in Australia (little penguin, Eudyptula minor ), in Antarctica (Adelie penguin, Pygoscelis adeliae ; emperor penguin, Aptenodytes forsteri ), and in subantarctic islands (king penguin, Aptenodytes patagonicus ; macaroni penguin, Eudyptes chrysolophus ; gentoo penguin, Pygoscelis papua ). All species use only vocal cues to identify their partner, but in territorial species the nest is used as a meeting point. In large species, such as the king and the emperor penguins, which do not have a nest, the brooder carries the egg or the small chick on the feet, while the mate, and then the chick, has to be located in the noisy colony without any topographical cue. According to theory, to extract a signal from background calls, animals analyze either frequency bands or modulations (amplitude and frequency modulations) of the partners call. The first coding-decoding system, used by nesting penguins, is easy to produce but costly in terms of analysis time. The second one, used by nonnesting penguins, is a vocal signature which is fast to analyze but costly to produce. This acoustic signal is particularly efficient as a means to locate immediately the partner on the move in a noisy crowd. Briefly, frequency analysis is enough to solve the relatively easy problem of individual recognition in nesting birds, while the complex temporal analysis of modulations of the two nonnesting penguins is an adaptation to extreme acoustic and breeding conditions. The macaroni penguin, which we have begun to test, seems to use both a frequency code similar to that of the other nesting species and a temporal code close to the one of a nonnesting penguin species, but much simpler.

Cocktail-party effect in king penguin colonies

The king penguin, Aptenodytes patagonicus, breeds without a nest in colonies of several thousands of birds. To be fed, the chick must recognize the parents in a particularly noisy environment using only vocal cues. The call an adult makes when seeking the chick is emitted at a high amplitude level. Nevertheless, it is transmitted in a colonial context involving the noise generated by the colony and the screening effect of the bodies, both factors reducing the signal–to–noise ratio. In addition, the adult call is masked by a background noise with similar amplitude and spectral and temporal characteristics, enhancing the difficulty for the chick in finding its parents. We calculate that the maximum distance from the caller at which its signal can be differentiated from the background noise (signal–to–noise ratio equal to 1) should not exceed 8–9 m in a feeding area. But our tests show that, in fact, chicks can discriminate between the parental call and calls from other adults at a greater distance, even when call intensity is well below that of the noise of simultaneous calls produced by other adults. This capacity to perceive and extract the call of the parent from the ambient noise and particularly from the calls of other adults, termed the ‘cocktail–party effect’ in speech intelligibility tests, enhances the chicks ability to find its parents.

How do king penguins (Aptenodytes patagonicus apply the mathematical theory of information to communicate in windy conditions

In the king penguin (Aptenodytes patagonicus), both pair members alternate in incubating and rearing their chick. Mates can recognize each other among thousands of other birds in the hubbub of the colony using only acoustic signalling: the display call. Large penguin colonies are found on sub–Antarctic islands where strong winds blow throughout the year. We have shown by experiments under natural conditions that the level of background noise increases in windy conditions and thus leads to a diminution of the signal–to–noise ratio. Moreover the emergence level of the signal revealed by entropy calculation is statistically weaker in windy conditions. To achieve breeding success, birds must continue communicating in spite of the significant decrease in the total amount of information that can be transmitted in windy situations. For the first time, to our knowledge, we have shown that a bird species takes into account the constraints imposed by wind on their acoustic communication. In windy conditions, birds try to maintain the efficiency of communication by increasing both the number of calls emitted and the number of syllables per call. This result conforms with predictions from the mathematical theory of communication: increased redundancy in a signal improves the probability of receiving a message in a noisy channel.

Penguins use the two-voice system to recognize each other

The sound–producing structure in birds is the syrinx, which is usually a two–part organ located at the junction of the bronchi. As each branch of the syrinx produces sound independently, many birds have two acoustic sources. Thirty years ago, we had anatomical, physiological and acoustical evidence of this twovoice phenomenon but no function was known. In songbirds, often these two voices with their respective harmonics are not activated simultaneously but they are obvious in large penguins and generate a beat pattern which varies between individuals. The emperor penguin breeds during the Antarctic winter, incubating and carrying its egg on its feet. Without the topographical cue of a nest, birds identify each other only by vocal means when switching duties during incubation or chick rearing. To test whether the twovoice system contains the identity code, we played back the modified call of their mate to both adults and also the modified call of their parents to chicks. Both the adults and the chicks replied to controls (two voices) but not to modified signals (one voice being experimentally suppressed). Our experiments demonstrate that the beat generated by the interaction of these two fundamental frequencies conveys information about individual identity and also propagates well through obstacles, being robust to sound degradation through the medium of bodies in a penguin colony. The two–voice structure is also clear in the call of other birds such as the king penguin, another non–nesting species, but not in the 14 other nesting penguins. We concluded that the two–voice phenomenon functions as an individual recognition system in species using few if any landmarks to meet. In penguins, this coding process, increasing the call complexity and resisting sound degradation, has evolved in parallel with the loss of territoriality.

Adult Male Mice Emit Context-Specific Ultrasonic Vocalizations That Are Modulated by Prior Isolation or Group Rearing Environment

Social interactions in mice are frequently analysed in genetically modified strains in order to get insight of disorders affecting social interactions such as autism spectrum disorders. Different types of social interactions have been described, mostly between females and pups, and between adult males and females. However, we recently showed that social interactions between adult males could also encompass cognitive and motivational features. During social interactions, rodents emit ultrasonic vocalizations (USVs), but it remains unknown if call types are differently used depending of the context and if they are correlated with motivational state. Here, we recorded the calls of adult C57BL/6J male mice in various behavioral conditions, such as social interaction, novelty exploration and restraint stress. We introduced a modulator for the motivational state by comparing males maintained in isolation and males maintained in groups before the experiments. Male mice uttered USVs in all social and non-social situations, and even in a stressful restraint context. They nevertheless emitted the most important number of calls with the largest diversity of call types in social interactions, particularly when showing a high motivation for social contact. For mice maintained in social isolation, the number of calls recorded was positively correlated with the duration of social contacts, and most calls were uttered during contacts between the two mice. This correlation was not observed in mice maintained in groups. These results open the way for a deeper understanding and characterization of acoustic signals associated with social interactions. They can also help evaluating the role of motivational states in the emission of acoustic signals.

ON THE EXTRACTION OF SOME TIME DEPENDENT PARAMETERS OF AN ACOUSTIC SIGNAL BY MEANS OF THE ANALYTIC SIGNAL CONCEPT. ITS APPLICATION TO ANIMAL SOUND STUDY

ABSTRACT When studying the acoustic signals of animals, it is often necessary to obtain the precise time structure of the frequency. It is also useful to extract the envelope of a signal or to modify a signal to obtain its frequency modulated part. This can be achieved digitally using the analytic signal concept. We deal here with the method. Definition and computation are described, its application is discussed and the performances are compared to those of sonagraph and zero- crossing methods. This paper presents a method enabling precise AM and FM analysis of different animal vocalisations. The method has also been used for the purpose of synthesis by extracting the frequency modulated part of a signal. It is found to be a powerful means of modifying some parameters of a natural signal without altering other features.

Are bird song complexity and song sharing shaped by habitat structure? An information theory and statistical approach

In songbirds, song complexity and song sharing are features of prime importance for territorial defence and mate attraction. These aspects of song may be strongly influenced by changes in social environment caused by habitat fragmentation. We tested the hypothesis that habitat fragmentation induced by human activities influences song complexity and song sharing in the skylark, a songbird with a very large repertoire and whose population recently underwent a large decline. We applied powerful mathematical and statistical tools to assess and compare song complexity and song sharing patterns of syllables and sequences of syllables in two populations: a declining population in a fragmented habitat, in which breeding areas are separated from each other by unsuitable surroundings, and a stable population in a continuous habitat. Our results show that the structure of the habitat influences song sharing, but not song complexity. Neighbouring birds shared more syllables and sequences of syllables in the fragmented habitat than in the continuous one. Habitat fragmentation seems thus to have an effect on the composition of elements in songs, but not on the number and complexity of these elements, which may be a fixed feature of song peculiar to skylarks.

Acoustic recognition in macaroni penguins: an original signature system

Penguins use vocal signatures as cues to identify their kin in dense colonies. Experimental studies of four species in two genera have pointed out that vocal signatures depend on the breeding ecology of the birds. Penguins that have a meeting site for pair members and chicks (genus Pygoscelis), as in many seabirds, have a less complex vocal signature than penguins that have no nests (genus Aptenodytes). To investigate whether this pattern would extend to other nest-building penguins, we studied the vocal signature in a species of the genus Eudyptes, the macaroni penguin, E. chrysolophus. Temporal and spectral features of signatures were analysed to determine which were likely to encode individual identity. Using a methodology derived from the theory of information, we measured and compared the amount of information given by each of these parameters by means of a stereotypy index. We then tested their effective efficiency in playback experiments of modified calls. Like Aptenodytes species, the macaroni penguins used a double coding system that integrated information in both the temporal and spectral domains. The encoding was made through the tempo given by the successive syllables of the call and the harmonic content of the call. However, information was not complementary but was mostly repeated in both domains, and variables were unidimensional as in Pygoscelis signatures. These results point out an original and simple signature system in macaroni penguins. Although they support the hypothesis of simpler systems in nest-building species they also reveal more subtle differences within this category.