Song syllable discrimination by song sparrows (Melospiza melodia)

A habituation test paradigm was used to examine the responses of free-living territorial adult male song sparrows (Melospiza melodia) to a range of synthetic songs. The three-phrased test songs differed from one another in having either conspecific or heterospecific (swamp sparrow, M. georgiana) syllables, or silence, in the second phrase. Subjects were exposed to repeated presentations of one song type until their approach distance to a loudspeaker increased. In one experiment, birds were habituated with a song consisting of three phrases of song sparrow syllables and then tested for generalization to either novel song sparrow syllables in the second phrase, swamp sparrow syllables, or silence. Birds discriminated between song sparrow syllables on two response measures, and between song sparrow and swamp sparrow syllables on one measure. In a second experiment, after habituation to a song with swamp sparrow syllables in the second phrase, birds did not generalize to novel song sparrow syllables, but they did generalize to novel swamp sparrow syllables. Thus song sparrows make finer distinctions among conspecific syllable variants than with alien syllables. The results further suggest that subtle species-specific differences in note structure within syllables are discriminated by song sparrows and potentially provide an adequate basis for individual recognition by song.     

Song perception during the sensitive period of song learning in zebra finches (Taeniopygia guttata)

The sensitive period is a special time for auditory learning in songbirds. However, little is known about perception and discrimination of song during this period of development. The authors used a go/no-go operant task to compare discrimination of conspecific song from reversed song in juvenile and adult zebra finches (Taeniopygia guttata), and to test for possible developmental changes in perception of syllable structure and syllable syntax. In Experiment 1, there were no age or sex differences in the ability to learn the discrimination, and the birds discriminated the forward from reversed song primarily on the basis of local syllable structure. Similar results were found in Experiment 2 with juvenile birds reared in isolation from song. Experiment 3 found that juvenile zebra finches could discriminate songs on the basis of syllable order alone, although this discrimination was more difficult than one based on syllable structure. The results reveal well-developed song discrimination and song perception in juvenile zebra finches, even in birds with little experience with song.     

Amplitude modulation of sexy phrases is salient for song attractiveness in female canaries (Serinus canaria)

Song discrimination and recognition in songbird species have usually been studied by measuring responses to song playbacks. In female canaries, Serinus canaria, copulation solicitation displays (CSDs) are used as an index of female preferences, which are related to song recognition. Despite the fact that many studies underline the role of song syntax in this species, we observed that short segments of songs (a few seconds long) are enough for females to discriminate between conspecific and heterospecific songs, whereas such a short duration is not sufficient to identify the syntax rules. This suggests that other cues are salient for song recognition. In this experiment, we investigated the influence of amplitude modulation (AM) on the responses (CSDs) of female canaries to song playbacks. We used two groups of females: (1) raised in acoustic isolation and (2) raised in normal conditions. When adult, we tested their preferences for sexy phrases with different AMs. We broadcast three types of stimuli: (1) songs with natural canary AM, (2) songs with AM removed, or (3) song with wren Troglodytes troglodytes AM. Results indicate that female canaries prefer and have predispositions for a song type with the natural canary AM. Thus, this acoustic parameter is a salient cue for song attractiveness.     

Discrimination between host songs by brood parasitic brown-headed cowbirds ( Molothrus ater)

Songbirds can learn both to produce and to discriminate between different classes of acoustic stimuli. Varying levels of auditory discrimination may improve the fitness of individuals in certain ecological and social contexts and, thus, selection is expected to mold the cognitive abilities of different species according to the potential benefits of acoustic processing. Although fine-scale auditory discrimination of conspecific songs and calls has been frequently reported for brood parasitic brown-headed cowbirds (Molothrus ater), it remains unclear why and how they perceive differently the songs of their many host species. Using habituation-dishabituation paradigms and measuring behavioral and physiological (heart-rate) responses, we found that captive female cowbirds consistently discriminated between songs of two host species, the song sparrow (Melospiza melodia) and the red-winged blackbird (Agelaius phoeniceus). Playback experiments with stimuli composed of con-specific followed by heterospecific vocalizations in the field also demonstrated discrimination between these heterospecific songs even though cowbirds were not attracted to playbacks of either host species' songs alone. Our results do not directly support a nest-searching function of heterospecific song discrimination by cowbirds and are most consistent with a function of the parasites' avoidance of attacks by their agressive hosts. These data demonstrate discrimination between heterospecific vocalizations by brown-headed cowbirds and add a novel dimension to the already expansive auditory perceptual abilities of brood parasitic species and other songbirds. Electronic Publication     

Distributed recognition of natural songs by European starlings

Individual vocal recognition behaviors in songbirds provide an excellent framework for the investigation of comparative psychological and neurobiological mechanisms that support the perception and cognition of complex acoustic communication signals. To this end, the complex songs of European starlings have been studied extensively. Yet, several basic parameters of starling individual vocal recognition have not been assessed. Here we investigate the temporal extent of song information acquired by starlings during vocal recognition learning. We trained two groups of starlings using standard operant conditioning techniques to recognize several songs from two conspecific male singers. In the first experiment we tested their ability to maintain accurate recognition when presented with (1) random sequences of 1–12 motifs (stereotyped song components) drawn from the training songs, and (2) 0.1–12-s excerpts of continuous song drawn from the training songs. We found that song recognition improved monotonically as more vocal material is provided. In the second experiment, we systematically substituted continuous, varying length regions of white noise for portions of the training songs and again examined recognition accuracy. Recognition remained above chance levels for all noise substitutions tested (up to 91% of the training stimulus) although all but the smallest substitutions led to some decrement in song recognition. Overall, above chance recognition could be obtained with surprisingly few motifs, short excerpts of song, and in the absence of large portions of the training songs. These results suggest that starlings acquire a representation of song during individual vocal recognition learning that is robust to perturbations and distributed broadly over large portions of these complex acoustic sequences.     

Auditory scene analysis in estrildid finches (Taeniopygia guttata and Lonchura striata domestica): a species advantage for detection of conspecific song

Operant-conditioning techniques were used to investigate the ability of zebra finches (Taeniopygia guttata) and Bengalese finches (Lonchura striata domestica) to detect a zebra finch or a Bengalese finch target song intermixed with other birdsongs. Sixteen birds were trained to respond to the presence of a particular target song, either of their own species (n = 8) or of another species (n = 8). The birds were able to learn a discrimination between song mixtures that contained a target song and song mixtures that did not, and they were able to maintain their response to the target song when it was mixed with novel songs. Zebra finches, but not Bengalese finches, learned the discrimination with a conspecific target more quickly and were worse at detecting a Bengalese finch in the presence of a conspecific song. The results indicate that selective attention to birdsongs within an auditory scene is related to their biological relevance.     

Cross-fostering diminishes song discrimination in zebra finches (Taeniopygia guttata)

Song-production, -discrimination, and -preferences in oscine birds are dually influenced by species identity and the ontogenetic environment. The cross-fostering of a model species for recognition research, the zebra finch (Taeniopygia guttata) into heterospecific nests of the Bengalese finch (Lonchura striata vars. domestica) allows an exploration of the sensory limits of early development and the effects of species-specific acoustic cues upon song discrimination in adulthood. To quantify the song preferences of female and male normal-reared (control) and Bengalese finch fostered zebra finches, we recorded multiple behavioral measures, including spatial proximity, vocalization rates and response latency, during sequential song-playback choice-trials using both tutor species’ songs and the songs of two other ecologically relevant Australian species, the owl finch (Taeniopygia bichenovii) and the star finch (Neochmia ruficauda). Response strength was variable between the different measures, but no differences were detected within the specific behavioral responses towards the song playbacks of the two sexes. Control subjects strongly preferred their own species’ songs while Bengalese-fostered zebra finches exhibited reduced song discrimination between con-, tutor-, and heterospecific songs. Overall behavioral responsiveness was also modulated by social ontogeny. These results indicate a difference in the strength of preference for song that is dependent on the species identity of the rearing environment in oscine birds and illustrate the role of multiple behavioral measures and ecologically relevant stimulus species selection in behavioral research using zebra finches.     

Integration of melody and text in memory for songs

Two experiments examined whether the memory representation for songs consists of independent or integrated components (melody and text). Subjects heard a serial presentation of excerpts from largely unfamiliar folksongs, followed by a recognition test. The test required subjects to recognize songs, melodies, or texts and consisted of five types of items: (a) exact songs heard in the presentation; (b) new songs; (c) old tunes with new words; (d) new tunes with old words; and (e) old tunes with old words of a different song from the same presentation (‘mismatch songs’). Experiment 1 supported the integration hypothesis: Subjects' recognition of components was higher in exact songs (a) than in songs with familiar but mismatched components (e). Melody recognition, in particular, was near chance unless the original words were present. Experiment 2 showed that this integration of melody and text occurred also across different performance renditions of a song and that it could not be eliminated by voluntary attention to the melody.     

The organisation of musical semantic memory: Evidence from false memories for familiar songs

By adapting a well-known paradigm for studying memory for words—the Deese-Roediger-McDermott or DRM paradigm (Deese, 1959, Roediger & McDermott, 1995)—the two experiments reported here explore memory for song titles and song clips. Participants were presented with five song titles (Experiment 1a) or five 30-second song clips (Experiment 1b) for each of nine popular artists (e.g., Robbie Williams). The most popular song identified for each artist in a pilot task was omitted from the sets of titles/clips. Following a distractor task, participants were asked to write down as many of the songs as they could recall. They were also asked to return a week later and complete a second recall task. Participants falsely recalled a significant number of the related but non-presented songs in both experiments and this increased a week later, while correct recall for presented items decreased. The results are discussed in terms of theory for musical memory as well as in the context of providing a novel method for exploring the organisation of musical memory.     

Vocal syntax development in the white-crowned sparrow (Zonotrichia leucophrys).

Birdsong development exemplifies the interplay between experience and predisposition that occurs during behavioral ontogeny. Songbirds must hear song models to develop normal song, yet they preferentially learn conspecific song when given a choice in the laboratory. To the extent that features guiding this selective learning are pre-encoded in the brain, such features should also develop in the song of young birds not exposed to them in tutor models. To investigate whether song syntax-phrase number and order--is such a feature in the white-crowned sparrow (Zonotrichia leucophrys), the authors tutored males of this species with separate phrase models. Birds learned and assembled these into songs of species-typical sequence, suggesting that syntax is to some degree pre-encoded in white-crowned sparrows. Birds also learned heterospecific phrases, confirming previous evidence that note phonology is not the primary cue for selective song learning in this species.     

Two-way interactions between music and language: Evidence from priming recognition of tune and lyrics in familiar songs

A priming technique was employed to study the relations between melody and lyrics in song memory. The procedure involved the auditory presentation of a prime and a target taken from the same song, or from unrelated but equally familiar songs. To promote access to memory representations of songs, we varied the format of primes and targets, which were either spoken or sung, using the syllable /1a/. In each of the four experiments, a prime taken from the same song as the target facilitated target recognition, independently of the format in which it occurred. The facilitation effects were also found in conditions close to masked priming because prime recognizability was very low, as assessed in Experiment 1 by d' measures. Above all, backward priming effects were observed in Experiments 2, 3, and 4, where song order was reversed in the prime-target sequence, suggesting that words and tones of songs are not connected by strict temporal contingencies. Rather, the results indicate that, in song memory, text and tune are related by tight connections that are bidirectional and automatically activated by relatively abstract information. Rhythmic similarity between linguistic stress pattern and musical meter might account for these priming effects.     

Direct social contacts override auditory information in the song-learning process in starlings (Sturnus vulgaris)

Social influence on song acquisition was studied in 3 groups of young European starlings raised under different social conditions but with the same auditory experience of adult song. Attentional focusing on preferred partners appears the most likely explanation for differences found in song acquisition in relation to experience, sex, and song categories. Thus, pair-isolated birds learned from each other and not from broadcast live songs, females did not learn from the adult male tutors, and sharing occurred more between socially associated peers. On the contrary, single-isolated birds clearly copied the adult songs that may have been the only source of attention stimulation. Therefore, social preference appears as both a motor for song learning and a potential obstacle for acquisition from nonpreferred partners, including adults.     

Behavioral consequences of song learning: Discrimination of song types by male white-crowned sparrows

The functional significance of learned population differences in male song in the white-crowned sparrow was explored in natural populations using playback tests. Laboratory results have shown that learning of the population-specific song seems to take place in early life and is strongly dependent upon the nature of the auditory experience at that time. However, the varied results of recent studies make it difficult to reach a confident conclusion about the ecological functions of song learning. The present research took advantage of naturally occurring variation in the differences between songs of adjacent populations to determine a function relating degree of difference in song to intensity of territorial singing elicited. Applying a typological evaluation of syllable structure to the four segments of the song allowed a crude quantitative ranking of the differences between local songs and playback stimuli. These results, together with those of other studies, suggest a unimodal aggressive response function of males to songs of other males. A maximum response to songs slightly different from the local song environment suggests that male exclusion based upon acquired song components may contribute to the maintenance of discrete and stable song dialects.     

Testing the absolute-tempo hypothesis: Context effects for familiar and unfamiliar songs

In two experiments, we investigated context effects on tempo judgments for familiar and unfamiliar songs performed by popular artists. In Experiment 1, participants made comparative tempo judgments to a remembered standard for song clips drawn from either a slow or a fast context, created by manipulating the tempos of the same songs. Although both familiar and unfamiliar songs showed significant shifts in their points of subjective equality toward the tempo context values, more-familiar songs showed significantly reduced contextual bias. In Experiment 2, tempo pleasantness ratings showed significant context effects in which the ordering of tempos on the pleasantness scale differed across contexts, with the most pleasant tempo shifting toward the contextual values, an assimilation of ideal points. Once again, these effects were significant but reduced for the more-familiar songs. The moderating effects of song familiarity support a weak version of the absolute-tempo hypothesis, in which long-term memory for tempo reduces but does not eliminate contextual effects. Thus, although both relative and absolute tempo information appear to be encoded in memory, the absolute representation may be subject to rapid revision by recently experienced tempo-altered versions of the same song.     

Influence of social conditions in song sharing in the adult canary

In songbirds, experience of social and environmental cues during a discrete period after birth may dramatically influence song learning. In the canary, the ability to learn new songs is assumed to persist throughout life. The aim of the present study was to investigate whether social context could guide changes in adult song. Three groups of canaries were kept in different social and temporal conditions. Results showed that the multiple hierarchical levels of the canary song structure were affected by social environment: songs of males housed together for 2 years were more similar than those of males that spent the same time in individual cages in regard to acoustic parameters, syllable repertoire and repertoire of sequences of two-syllable types. However, social housing did not result in the emergence of a group-specific vocal signature within songs. In conclusion, these results suggested that under the influence of social factors, a copying process could allow adult canaries to adjust, at least in part, their songs to those of other individuals.     

Perception of temporal properties in self-generated songs by Bengalese finches (Lonchura striata var. domestica)

In the songbird forebrain, neuronal selectivity for temporal properties of each bird's self-generated song has been well described, but the behavioral and perceptual correlates of this selectivity are not known. By operant procedures, the authors trained Bengalese finches (Lonchura striata var. domestica) to discriminate between songs that were played normally and in reverse. Male Bengalese finches learned the discrimination quicker when their self-generated song was used as a stimulus than when a song of another conspecific bird was used. When the global note order was retained but each note was locally reversed, the song was more likely to be regarded as a forward song by the singer himself, but not by other birds. These results provide psychophysical evidence that the special processing of the self-generated song observed at the neural level might reflect an individual's perception of his self-produced song.     

Discrimination of individual vocalizations by black-capped chickadees (Poecile atricapilla)

The auditory perceptual abilities of male black-capped chickadees (Poecile atricapilla) were examined using an operant go/no-go discrimination among 16 individual vocalizations recorded at 5 m. The birds learned to discriminate about equally well among eight male chickadee fee-bee songs and eight female zebra finch (Taeniopygia guttata) distance calls. These results do not indicate that chickadees have a species-specific advantage in individual recognition for conspecific over heterospecific vocalizations. We then transferred the chickadees to a discrimination of the same songs and calls rerecorded at a moderate distance. These results showed accurate transfer of discrimination from 16 vocalizations recorded at 5 m to novel versions of the same 16 songs and calls rerecorded at 25 m. That is, chickadees recognized individual songs and calls despite degradation produced by rerecording at 25 m. Identifying individual vocalizations despite their transformation by distance cues is here described as a biologically important example of perceptual constancy.     

Factors influencing song development in the white-crowned sparrow (Zonotrichia leucophrys)

Nestling white-crowned sparrows (Zonotrichia leucophrys nuttalli) were hand-reared in sound-isolation chambers under a variety of conditions. The songs of total isolates were compared with songs of birds tutored with song, and the number of inputs sufficient for a bird to produce a normal song was explored. The flexibility of the song learning system was investigated with a range of tape-recorded tutor songs: alien dialects, alien subspecies, alien species, alternating alien dialects, and an aberrant song. Adult songs were obtained for 40 males and 7 testosterone-injected females. All of the tutor songs could be learned. Also, some birds learned elements of an alien species' song. Birds tutored with two songs copied one or the other, were bilingual, or sang a hybrid of the two. No bird presented with fewer than 120 songs learned the tutor song; 2 birds tutored with 252 songs copied the tutor song. It is concluded that the song learning system is quite flexible, hat the results obtained with tape-tutors are very different from those with social tutors, and that there may be an interaction between total number of song inputs and the number presented on a single day. Some implications of these data for physiological mechanisms and the possible functional significance of the acquisition system are discussed.     

Song recognition in zebra finches: Are there sensitive periods for song memorization?

Male zebra finches learn to sing songs that they hear between 25 and 65 days of age, the sensitive period for song learning. In this experiment, male and female zebra finches were exposed to zebra finch songs either before (n = 9) or during (n = 4) the sensitive period. Following song exposure, recognition memory for the songs was assessed with an operant discrimination between familiar and novel songs. Zebra finches that were exposed to songs between 22 and 30 days of age discriminated between familiar and novel songs; zebra finches exposed to songs from 9 to 17 days of age did not. Failure to memorize songs heard prior to the sensitive period may contribute to the exclusion of those songs from the repertoire of songbirds.     

Timbre control in zebra finch (Taeniopygia guttata) song syllables

Zebra finch (Taeniopygia guttata) song syllables often include harmonically related frequency components. These harmonics may be suppressed, and this differential emphasis varies between the syllables in a song and between individual birds' songs. These patterns of harmonic suppression are timbre. Individual syllables' patterns of harmonic suppression are constant with adult males' songs. Young males that imitate the songs of older males also imitate their patterns of harmonic suppression. Syringeal denervation distorts these patterns, which suggests that they are produced through active control of the vocal organ. The selective suppression and emphasis of some harmonics creates a great number of possible timbre variants for any one syllable. These add signal diversity to the limited array of frequency modulations and range of fundamental frequencies found in zebra finch song. Analyses of bird song that disregard timbre may overlook a feature that is important in vocal communication.