Auditory pattern discrimination in budgerigars (Melopsittacus undulatus).

Auditory patterns carry information in human speech at multiple levels, including the surface relationships between sounds within words in phonology and the abstract structures of syntax. The sequences of other animal vocalizations, such as birdsong, can also be described as auditory patterns, but few studies have probed how the sequences are perceived at multiple levels. Past work shows that a small parrot species, the budgerigar (Melopsittacus undulatus), exceeds other birds in sequence perception and is even sensitive to abstract structure. But it is not known what level of auditory analysis is dominant in perception or what limits might exist in sensitivity to abstract structure. Here, budgerigars were tested on their ability to discriminate changes in an auditory pattern, AAB, i.e. sound-same different, to ask how they attended to surface relationships among the sounds and the abstract relationships of same/different among the elements. The results show that the budgerigars primarily used surface transitions between the sounds when discriminating the sequences, but were able to use the abstract relationships to a limited extent, largely restricted to two elements. This study provides insight into how budgerigars extract information from conspecific vocalizations and how their capacities compare to human speech perception.

Discrimination of natural acoustic variation in vocal signals.

Studies of acoustic communication often focus on the categories and units of vocalizations, but subtle variation also occurs in how these signals are uttered. In human speech, it is not only phonemes and words that carry information but also the timbre, intonation, and stress of how speech sounds are delivered (often referred to as "paralinguistic content"). In non-human animals, variation across utterances of vocal signals also carries behaviorally relevant information across taxa. However, the discriminability of these cues has been rarely tested in a psychophysical paradigm. Here, we focus on acoustic communication in the zebra finch (Taeniopygia guttata), a songbird species in which the male produces a single stereotyped motif repeatedly in song bouts. These motif renditions, like the song repetitions of many birds, sound very similar to the casual human listener. In this study, we show that zebra finches can easily discriminate between the renditions, even at the level of single song syllables, much as humans can discriminate renditions of speech sounds. These results support the notion that sensitivity to fine acoustic details may be a primary channel of information in zebra finch song, as well as a shared, foundational property of vocal communication systems across species.

What can animal communication teach us about human language?

Language has been considered by many to be uniquely human. Numerous theories for how it evolved have been proposed but rarely tested. The articles in this theme issue consider the extent to which aspects of language, such as vocal learning, phonology, syntax, semantics, intentionality, cognition and neurobiological adaptations, are shared with other animals. By adopting a comparative approach, insights into the mechanisms and origins of human language can be gained. While points of agreement exist among the authors, conflicting viewpoints are expressed on several issues, such as the presence of proto-syntax in animal communication, the neural basis of the Merge operation, and the neurogenetic changes necessary for vocal learning. Future comparative research in animal communication has the potential to teach us even more about the evolution, neurobiology and cognitive basis of human language. This article is part of the theme issue 'What can animal communication teach us about human language?'

Sound sequences in birdsong: how much do birds really care?

The complex and melodic nature of many birds' songs has raised interest in potential parallels between avian vocal sequences and human speech. The similarities between birdsong and speech in production and learning are well established, but surprisingly little is known about how birds perceive song sequences. One popular laboratory songbird, the zebra finch (Taeniopygia guttata), has recently attracted attention as an avian model for human speech, in part because the male learns to produce the individual elements in its song motif in a fixed sequence. But psychoacoustic evidence shows that adult zebra finches are relatively insensitive to the sequential features of song syllables. Instead, zebra finches and other birds seem to be exquisitely sensitive to the acoustic details of individual syllables to a degree that is beyond human hearing capacity. Based on these findings, we present a finite-state model of zebra finch perception of song syllable sequences and discuss the rich informational capacity of their vocal system. Furthermore, we highlight the abilities of budgerigars (Melopsittacus undulatus), a parrot species, to hear sequential features better than zebra finches and suggest that neurophysiological investigations comparing these species could prove fruitful for uncovering neural mechanisms for auditory sequence perception in human speech. This article is part of the theme issue 'What can animal communication teach us about human language?'

Strain differences in hearing in song canaries.

Belgian Waterslager song canaries, bred for hundreds of years for a low-pitched song, have also acquired an inherited high-frequency hearing loss associated with hair cell abnormalities. Here, auditory thresholds measured using auditory brainstem responses and psychophysical methods in three different strains of canaries are compared: Belgian Waterslagers, American Singers, and Borders. Border canaries have not been bred for song characteristics while American Singer canaries have been bred for song only since the 1930s. Results show that American Singer canaries also have elevated high frequency thresholds that are similar to those of the Belgian Waterslager, while Border canaries have normal thresholds. These results strengthen the case that song canary breeders in selecting for song characteristics may have inadvertently selected for hearing abnormalities.

How canaries listen to their song: Species-specific shape of auditory perception.

The melodic, rolling songs of canaries have entertained humans for centuries and have been studied for decades by researchers interested in vocal learning, but relatively little is known about how the birds listen to their songs. Here, it is investigated how discriminable the general acoustic features of conspecific songs are to canaries, and their discrimination abilities are compared with a small parrot species, the budgerigar. Past experiments have shown that female canaries are more sexually responsive to a particular song element-the "special" syllables-and consistent with those observations, it was found that special syllables are perceptually distinctive for canaries. It is also shown that canaries discriminate the subtle differences among syllables and phrases using spectral, envelope, and temporal fine structure cues. Yet, while canaries can hear these fine details of the acoustic structure of their song, the evidence overall suggests that they listen at a more global, phrase by phrase level, rather than an analytic, syllable by syllable level, except when attending to some features of special syllables. These results depict the species-specific shape of auditory perception in canaries and lay the groundwork for future studies examining how song perception changes seasonally and according to hormonal state.

Dynamic sex-specific responses to synthetic songs in a duetting suboscine passerine.

Many bird species produce temporally coordinated duets and choruses, requiring the rapid integration of auditory perception and motor production. While males and females of some species are known to participate in these displays for sex-specific purposes, few studies have identified perceptual features that trigger sex-specific contributions of coordinated song. Especially little is known about perception and production in duetting suboscine passerines, which are thought to have innate songs and largely static, rather than dynamic, vocal behavior. Here, we used synthetic stimuli in a playback experiment on chestnut-backed antbirds (Myrmeciza exsul) to (1) test whether differences in song frequency (Hz) can trigger sex-specific vocal behavior in a suboscine passerine (2) test for the functions of duetting in males and females of this species, and (3) determine whether these suboscines can dynamically adjust the temporal and spectral features of their songs. We found sex-specific responses to synthetic playback manipulated in song frequency (Hz), providing evidence that in this context males sing in duets for general territory defense and females join in for mate guarding purposes. In addition, we found that the birds altered the frequency, duration, and timing of their songs depending on the frequency of the playback songs. Thus, we show that these birds integrate spectral and temporal information about conspecific songs and actively modulate their responses in sex-specific ways.

Relative salience of syllable structure and syllable order in zebra finch song.

There is a rich history of behavioral and neurobiological research focused on the 'syntax' of birdsong as a model for human language and complex auditory perception. Zebra finches are one of the most widely studied songbird species in this area of investigation. As they produce song syllables in a fixed sequence, it is reasonable to assume that adult zebra finches are also sensitive to the order of syllables within their song; however, results from electrophysiological and behavioral studies provide somewhat mixed evidence on exactly how sensitive zebra finches are to syllable order as compared, say, to syllable structure. Here, we investigate how well adult zebra finches can discriminate changes in syllable order relative to changes in syllable structure in their natural song motifs. In addition, we identify a possible role for experience in enhancing sensitivity to syllable order. We found that both male and female adult zebra finches are surprisingly poor at discriminating changes to the order of syllables within their species-specific song motifs, but are extraordinarily good at discriminating changes to syllable structure (i.e., reversals) in specific syllables. Direct experience or familiarity with a song, either using the bird's own song (BOS) or the song of a flock mate as the test stimulus, improved both male and female zebra finches' sensitivity to syllable order. However, even with experience, birds remained much more sensitive to structural changes in syllables. These results help to clarify some of the ambiguities from the literature on the discriminability of changes in syllable order in zebra finches, provide potential insight on the ethological significance of zebra finch song features, and suggest new avenues of investigation in using zebra finches as animal models for sequential sound processing.