Reversal of hypoglossal dominance in canaries following unilateral syringeal denervation

Summary The song of intact male canaries develops under the almost exclusive control of the left hypoglossus and left syringeal half. Section of the left hypoglossus or of its tracheosyringealis branch induces the right hypoglossus to assume a dominant control over vocal behavior. When this operation is done during the first two weeks after hatching the ensuing song is under sole right hypoglossal control; if done during the third and fourth week after hatching song develops under shared right-left hypoglossal control. In either case the quality of song is close to that of intact birds of comparable age. If the left hypoglossal innervation to the syrinx is cut when song development is already well under way (plastic song) or after stable adult song has been acquired, then the quality of song developed is markedly poorer than that of controls. From these observations we may conclude that: 1) Left or right hypoglossal dominance are not necessary for the production of normal canary song. 2) Soon after hatching either hypoglossus has the potential to assume a dominant role in song development. 3) The ability of the right hypoglossus to develop normal song decreases as birds master song under dominant left hypoglossal control. It is suggested that hypoglossal dominance and dominance reversal may provide convenient material to study neuronal changes related to the learning of new motor tasks.Abbreviations e.l. external labium - i.t.m. internal tympaniform membrane - ts tracheosyringealis Steven Green, Myron C. Baker and Timothy DeVoogd provided invaluable help on statistical analysis. Steven Green also read the text and made useful suggestions.     

Learning-Related Neuronal Activation in the Zebra Finch Song System Nucleus HVC in Response to the Bird's Own Song

Like many other songbird species, male zebra finches learn their song from a tutor early in life. Song learning in birds has strong parallels with speech acquisition in human infants at both the behavioral and neural levels. Forebrain nuclei in the 'song system' are important for the sensorimotor acquisition and production of song, while caudomedial pallial brain regions outside the song system are thought to contain the neural substrate of tutor song memory. Here, we exposed three groups of adult zebra finch males to either tutor song, to their own song, or to novel conspecific song. Expression of the immediate early gene protein product Zenk was measured in the song system nuclei HVC, robust nucleus of the arcopallium (RA) and Area X. There were no significant differences in overall Zenk expression between the three groups. However, Zenk expression in the HVC was significantly positively correlated with the strength of song learning only in the group that was exposed to the bird's own song, not in the other two groups. These results suggest that the song system nucleus HVC may contain a neural representation of a memory of the bird's own song. Such a representation may be formed during juvenile song learning and guide the bird's vocal output.     

Neural song preference during vocal learning in the zebra finch depends on age and state

The zebra finch acquires its song by first memorizing a model song from a tutor and then matching its own vocalizations to the memory trace of the tutor song, called a template. Neural mechanisms underlying this process require a link between the neural memory trace and the premotor song circuitry, which drives singing. We now report that a premotor song nucleus responds more to the tutor song model than to every other stimulus examined, including the bird's own song (BOS). Neural tuning to the song model occurred only during waking and peaked during the template-matching period of development, when the vocal motor output is sculpted to match the tutor song. During the same developmental phase, the BOS was the most effective excitatory stimulus during sleep. The preference for BOS compared to tutor song inverted with sleep/wake state. Thus, song preference shifts with development and state.     

The Development of Geographic Song Preferences in Female Song Sparrows Melospiza melodia

Previous work suggests that early learning plays a role in auditory preferences of female songbirds. We explored whether early experience shapes preferences for local geographic song in female song sparrows ( Melospiza melodia ), a species that prefers local geographic song in adulthood. We hand-reared females from two locations 450 km apart and controlled song exposure early in life. To examine the effects of experience and inherited factors on geographic song preferences, we used a counterbalanced design. Females from each location were tutored with either natal song (recorded from their location of capture) or non-natal song (recorded from the other location) and their preferences for tutor vs. non-tutor song assessed in adulthood. We also examined song preferences in isolate females reared with no song experience. We found that tutored females preferred their tutor song over non-tutor song, regardless of their capture location. Furthermore, birds not exposed to song (isolates) had no geographic song preferences. Thus, song exposure early in life clearly plays an important role in adult female preferences for local song in this species.     

Materialism and Sensuality: Visualizing the Devil in the Festival of Our Lady of Urkupiña

During the Festival of Our Lady of Urkupin ¨ a, held every August in Quillacollo, Bolivia, an estimated half million pilgrims petition Our Lady for wealth and material goods. The festival has become a national event and now includes the participation of government officials, including the President, as well as widespread coverage by the media. The festival has also become a point of controversy and contestation over the meanings of Bolivian nationalism, commercialism, and economic development. These issues, as I will show, have been articulated through criticisms of the festival as being too sensuous and overly commercialized. Through an examination of media coverage of the festival of Urkupiña, this article focuses on the ways that "evils" of materialism and sensuality are represented visually. The Bolivian state and the Catholic Church have sought to define the meaning of the festival in terms of devotion and national identity while criticizing some festival practices associated with gaining wealth. Here I show how the folkloric and devotional elements of the festival are embodied in the dancers while the visual referent for the materialism of the festival is street vendors. I also discuss how the festival transforms the meanings of the ch'alla and other Andeanpractices that anthropologists have traditionally associated with resistance to capitalism (as found for example in the work of June Nash and Michael Taussig). The article concludes by noting that, rather than mapping these meanings ontogeneralized resistance, anthropologists must be prepared fully to engage contestations over devotion, material gain, sensuousness, and economic development. In short, we should also be prepared to dance with the devil.     

Simplified song in an island population of the bush warbler Cettia diphone

Song complexity and the repertoire of the bush warbler Cettia diphone were studied in an island and a mainland population in Japan. The song complexity (number of modulations in a song) was lower in the island population than in the mainland one. On the other hand, the repertoire size (number of song types per male) was larger on the island. Founder effect probably does not influence the island song because colonization occurred a very long time ago. Sound transmission properties of the habitat and weak selection pressure for interspecific identification might have influenced the island song, but these cannot elucidate the simpler song on the island. There is a possibility that social conditions affect song complexity in the populations; the highly polygynous mating system of the mainland population yields strong selection pressure toward complex song through acquisition of mates and/or establishment of quality territories. On the other hand, variable songs on the island seem to be affected by cultural mutation. Received: September 3, 1999 / Accepted: December 9, 1999     

Non-breeding song rate reflects nutritional condition rather than body condition

Numerous studies have focused on song in songbirds as a signal involved in mate choice and intrasexual competition. It is expected that song traits such as song rate reflect individual quality by being dependent on energetic state or condition. While seasonal variation in bird song (i.e., breeding versus non-breeding song) and its neural substrate have received a fair amount of attention, the function and information content of song outside the breeding season is generally much less understood. Furthermore, typically only measures of condition involving body mass are examined with respect to song rate. Studies investigating a potential relationship between song rate and other indicators of condition, such as physiological measures of nutritional condition, are scant. In this study, we examined whether non-breeding song rate in male European starlings (Sturnus vulgaris) reflects plasma metabolite levels (high-density lipoproteins (HDL), albumin, triglycerides and cholesterol) and/or body mass. Song rate was significantly positively related to a principal component representing primarily HDL, albumin and cholesterol (and to a lesser degree plasma triglyceride levels). There was only a trend toward a significant positive correlation between song rate and body mass, and no significant correlation between body mass and the abovementioned principal component. Therefore, our results indicate that nutritional condition and body mass represent different aspects of condition, and that song rate reflects nutritional rather than body condition. Additionally, we also found that intra-individual song rate consistency (though not song rate itself) was significantly positively related to lutein levels, but not to body mass or nutritional condition. Together our results suggest that the relation between physiological measures of nutritional condition and song rate, as well as other signals, may present an interesting line of future research, both inside and outside the breeding season.     

Levels of Organization in the Song of the Bobolink (Icteridae: Dolichonyginae)

By comparison with other icterids, the bobolink (Dolichonyx oryzivorus) sings an unusually long and complex song. The songs of four male bobolinks were analysed in terms of units at four different levels of analysis: the figure, the figure-sequence, the song-pattern, and the song. The different levels of analysis show different degrees of stereotypy and variation. Individual males in the study area differed in the frequency of use of the different song-patterns and in the way they assembled them into songs. Immediate repetition of song patterns within a song was unusual. Songs sung from fixed perches were shorter than songs sung in flight. The results suggest that the length of bobolink song is in part a consequence of its use of the song as a part of an elaborate flight display, and that the unit we called the “song-pattern” was closest to the unit designated “song” in most other species.     

Individual,population, and geographic differentiation in advertising song of the Blyth’s reed warbler, <Emphasis Type="Italic">Acrocephalus dumetorum</Emphasis> (Sylvidae)

Processes of differentiation in the advertising song of the Blyth’s reed warbler have been studied both within local populations and over the species range. Individual vocal repertoires of males from the Moscow and Kostroma regions and the Southern Urals include 30 to 104 (on average, 55.3 ± 17.0) song types, with 45.9–84.4% (61.3 ± 10.5%) of song types being common to all males of a given population. The rate of accumulation of new common song types drops with an increase in the number of males included in comparison. The appearance of many song types in the repertoire cannot be attributed to imitation, at least within one geographic population. The repertoires of males from one geographic population are more similar to each other than to those from other populations. The degree of acoustic similarity of song repertoires between different populations is inversely related to the distance between them, but no such relationship has been revealed for the repertoires of individual males within the same population. Many song types are widespread almost throughout the species range. Songs of males from the southern part of the range differ in an accelerated performance rhythm, chaotic syntax, and unstable phonetic structure. Consideration is given to probable causes of song specificity in southern populations and to some parameters of individual song variation and other factors than can contribute to the formation of vocal repertoires in the Blyth’s reed warbler, such as nesting site fidelity, vocal mimicry, and song improvisation.     

Changes in Song Complexity Correspond to Periods of Female Fertility in Blue Grosbeaks (Guiraca caerulea)

Song complexity is thought to be a sexually selected trait in passerine birds; however, quantifying relevant parameters of song complexity is the first step in testing the theory that song complexity is a sexually selected trait. We show here that blue grosbeak (Guiraca caerulea) males sing a single song type but the properties of that song type vary between renditions. This pattern of song delivery potentially provides females with an opportunity to assess dimensions of song complexity other then repertoire size. Here we characterize song complexity using four measures: (i) element repertoire size, (ii) proportion of distinct song variant, (iii) song versatility, and (iv) syntax consistency. We studied the functional significance of song complexity by comparing measures of song complexity before and after periods of female fertility. We found that male blue grosbeaks sing more song variants, use more versatile arrangements of elements, and maintain more syntax consistency during the fertile period of their social mate than during their social mate's non‐fertile period. These results point to a functional relationship between changes in song complexity and periods of female fertility in this species.     

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.     

Neural auditory selectivity develops in parallel with song

The zebra finch learns his song by memorizing a tutor's vocalization and then using auditory feedback to match his current vocalization to this memory, or template. The neural song system of adult and young birds responds to auditory stimuli, and exhibits selective tuning to the bird's own song (BOS). We have directly examined the development of neural tuning in the song motor system. We measured song system responses to vocalizations produced at various ages during sleep. We now report that the auditory response of the song motor system and motor output are linked early in song development. During sleep, playback of the current BOS induced a response in the song nucleus HVC during the song practice period, even when the song consisted of little more than repeated begging calls. Halfway through the sensorimotor period when the song was not yet in its final form, the response to BOS already exceeded that to all other auditory stimuli tested. Moreover, responses to previous, plastic versions of BOS decayed over time. This indicates that selective tuning to BOS mirrors the vocalization that the bird is currently producing.     

Immune challenge mediates vocal communication in a passerine bird: an experiment

Secondary sexual characters may have evolved in part to signalresistance to parasites. Avian song has been hypothesized tobe involved in this process, but the role of parasites in modulatingacoustic communication systems in birds remains largely unknown,owing to lack of experiments. We studied the relationship betweenparasitism, testosterone, song performance, and mating successin male collared flycatchers (Ficedula albicollis) by experimentallychallenging their immune system with a novel antigen. We predictedthat a challenge of the immune system would reduce song performance,and that this reduction would be conditional on the size ofa visual sexual signal, the forehead patch that was previouslyfound to reflect resistance. An antagonistic linkage betweentestosterone and immune function would predict that a challengeof the immune system should suppress testosterone level. Animmunological treatment by sheep red blood cells (SRBCs) triggereda decrease in body mass, testosterone level, and song rate,but other song traits were not significantly affected by theantigen challenge. Initial testosterone level was associatedwith forehead patch size and all song traits except song rate.SRBC injection caused stronger reduction in song rate amongmales with smaller forehead patches, and the change in songrate was also predictable by song features such as strophe complexityand length. We show that song rate and other song characteristicsmay be important cues in male-male competition and female choice.These results suggest that parasite-mediated sexual selectionhas contributed in shaping a complex acoustic communicationsystem in the collared flycatcher, and that testosterone mayplay an important role in this process. Parasitism may drivea multiple signaling mechanism involving acoustic and visualtraits with different signal function.     

Song learning and mate choice in estrildid finches raised by two species

Male zebra finches, Taeniopygia guttata, and Bengalese finches, Lonchura striata, were raised normally or cross-fostered to the other species until 35 days of age and then housed until 70 days old with two song tutors, one from each species. Males raised normally based their song on that of the conspecific tutor whereas cross-fostered males also learned from the other tutor, reflecting both own-species bias and a tendency to learn from the tutor whose song was similar to the father's. Males raised by one bird from each species learned from the father as well as the tutor(s) but were more likely to learn from the tutor whose song was similar to the father's if the father was a conspecific. Comparing these song learning results with those from male choice tests suggests there is no direct link between song learning and sexual imprinting.     

Behavioural response to song and genetic divergence in two subspecies of white‐crowned sparrows (Zonotrichia leucophrys)

Divergence in sexual signals may drive reproductive isolation between lineages, but behavioural barriers can weaken in contact zones. Here, we investigate the role of song as a behavioural and genetic barrier in a contact zone between two subspecies of white‐crowned sparrows (Zonotrichia leucophrys). We employed a reduced genomic data set to assess population structure and infer the history underlying divergence, gene flow and hybridization. We also measured divergence in song and tested behavioural responses to song using playback experiments within and outside the contact zone. We found that the subspecies form distinct genetic clusters, and demographic inference supported a model of secondary contact. Song phenotype, particularly length of the first note (a whistle), was a significant predictor of genetic subspecies identity and genetic distance along the hybrid zone, suggesting a close link between song and genetic divergence in this system. Individuals from both parental and admixed localities responded significantly more strongly to their own song than to the other subspecies song, supporting song as a behavioural barrier. Putative parental and admixed individuals were not significantly different in their strength of discrimination between own and other songs; however, individuals from admixed localities tended to discriminate less strongly, and this difference in discrimination strength was explained by song dissimilarity as well as genetic distance. Therefore, we find that song acts as a reproductive isolating mechanism that is potentially weakening in a contact zone between the subspecies. Our findings also support the hypothesis that intraspecific song variation can reduce gene flow between populations.     

The Role of Song Phonology and Syntax in Vocal Learning Preferences in the Song Sparrow,Melospiza melodia

Are young songbirds ready to learn virtually any song, or are they predisposed to learn songs of their own species? To explore this question tests were conducted on the equipotentiality of auditory song learning stimuli in the song sparrow. 23 males reared as nestlings were exposed to tape recordings of their own and other species songs in early life and subsequent song production was analyzed for imitations. Birds exposed to natural song sparrow songs, including their fathers', and equal numbers of swamp sparrow songs, strongly preferred conspecific songs. They neither favored nor eschewed paternal songs despite having had access to them for 6–10 days as nestlings. In three other experiments synthetic songs were used in which some properties were held constant and others were systematically varied. Birds were exposed to 1–4 segmented songs varying in phrase order, tempo and syllable number, each synthesized in two versions, one from conspecific and the other from heterospecific (swamp sparrow) song syllables. With one-segmented songs (alien syntax) subjects favored conspecific over heterospecific syllable songs. Heterospecific syllables were rendered more acceptable by incorporation into two-segmented trilled songs (more song sparrow-like syntax). Heterogeneous summation of phonological and syntactical cues appeared to occur. There was also evidence of interaction between phonology and syntax. When another phrase type, the note complex, was added, in three- and four-segmented songs, a preference for conspecific syllables reappeared. Heterospecific syllables may be more readily accepted as a trilled sequence than without repetition, as in a note-complex. When phrase structure within four-segmented songs was varied, birds favored patterns most like normal conspecific song. We conclude that there are innate learning preferences in the song sparrow, based on note and syllabic structure (phonology and syllabic syntax), and temporal organization of phrases (segmental syntax), differing from those of the closely related swamp sparrow, Melospiza georgiana, in which song syntax plays no role in learning preferences.     

The importance of calling song and courtship song in female mate choice in the variable field cricket

Male field crickets produce calling songs, courtship songs, tactile signals and chemical signals. Although calling songs are known to play an important role in female mate choice, the importance of the other signals in mate choice is poorly understood. In the variable field cricket, Gryllus lineaticeps, females select mates, in part, based on variation in male calling song. Females prefer higher chirp rates, a trait which is partially dependent on male nutrient intake, and females prefer longer chirp durations, a trait which appears to be independent of male nutrient intake. We tested whether females also have preferences based on variation in male courtship song, and whether the structure of male courtship song varies with nutrient intake. First, we reexamined female preference for calling song chirp rate. Then, we examined: (1) female preference based on courtship song chirp rate; (2) the relative importance of calling song and courtship song chirp rate; (3) the nutrition dependence of courtship song chirp rate; and (4) the correlation between calling song and courtship song chirp rate. As reported previously, females preferred higher calling song chirp rates, and in addition, preferred higher courtship song chirp rates. Females were more likely to switch from a speaker broadcasting more attractive calling song to a speaker broadcasting less attractive calling song when the attractive calling song was associated with an unattractive courtship song than when it was associated with an attractive courtship song. Preferences based on courtship song may thus cause females to alter the choices that they made based on calling song. Males that received greater nutrients did not produce higher courtship song chirp rates. There was no correlation between calling song and courtship song chirp rate. As a result, the two traits may provide information to females about different aspects of male quality. Copyright 2000 The Association for the Study of Animal Behaviour.     

Signalling through acoustic windows: nightingales avoid interspecific competition by short-term adjustment of song timing

The function of bird song is closely linked to sexual selection through female choice and male–male competition, and thus variation in communication success is likely to have major fitness consequences for a singing male. A crucial constraint on signal transmission is imposed by background noise, which may include songs from other species. I investigated whether singing nightingales (Luscinia megarhynchos) avoid temporal overlap with the songs of other bird species in a playback experiment. I analysed the temporal song patterns of six males, each of which were exposed to songs of other species. The nightingales significantly avoided overlapping their songs with the playback songs, and started singing preferentially during the silent intervals between the heterospecific songs. This timing of song onset produced a greater variability in pause duration compared to the nightingales’ undisturbed solo singing. These findings suggest that birds adjust their song timing to avoid acoustic interference on short temporal scales, and thus are able to improve the efficiency of acoustic communication in complex sonic environments. Moreover, the results indicate that temporal song patterns can be affected by the songs of other species, and thus such influences should be taken into account when studying bird song in the field.     

Male territorial aggression does not drive conformity to local vocal culture in a passerine bird

In many songbird species, young individuals learn songs from neighbors and then settle nearby, thus creating neighborhoods of conformity to local vocal culture. In some species, individuals appear to postpone song learning until after dispersal, possibly to facilitate conformity to local dialects. Despite decades of study, we still lack a consensus regarding the selective pressures driving this delayed song learning. Two common hypothetical benefits to conformity, and thus delayed song learning, are rooted in territorial interactions; individuals preferentially produce local song either to avoid detection as new arrivals (deceptive mimicry) or to be more effectively recognized as conspecific territory holders. The dickcissel (Spiza americana) is an ideal species in which to study these hypotheses. Males of this species appear to delay song learning until they arrive at their first adult territory, each individual sings a single song type, and conformity to the local song culture is high. Using playback, we contradicted both of the territorial hypotheses described above; male dickcissels did not respond differentially to local vs foreign song playback treatment. We are confident in this lack of difference because dickcissels clearly responded less strongly to a third treatment, neighbor song, than to the other two treatments, demonstrating sufficient power in our experimental design (and providing the first evidence of the dear‐enemy effect in dickcissels). Our results raise the question of why dickcissels respond equally aggressively to both local and foreign songs when other bird species often show reduced aggression toward foreign song. If reduced aggression to foreign song is not ubiquitous in species that achieve conformity through delayed learning, then selection from aggressive territorial interaction seems unlikely to be a general explanation for such delayed learning. Reduced aggression in response to foreign songs in other species may be due to reduced exposure to the stimulus of foreign song or to different cost‐benefit trade‐offs when responding to songs that deviate from the local average.     

Relative threat and recognition ability in the responses of tropical mockingbirds to song playback

It has been suggested that individual recognition based on song may be constrained by repertoire size in songbirds with very large song repertoires. This hypothesis has been difficult to test because there are few studies on species with very large repertoires and because traditional experiments based on the dear enemy effect do not provide evidence against recognition. The tropical mockingbird, Mimus gilvus, is a cooperative breeder with very large song repertoires and stable territorial neighbourhoods. The social system of this species allowed us to test individual recognition based on song independently from the dear enemy effect by evaluating male response to playback of strangers, neighbours (from shared and unshared boundaries), co-males (i.e. other males in the same social group) and own songs. Although subjects did not show a dear enemy effect, they were less aggressive to co-males than to all other singers. Our results suggest that recognition in tropical mockingbirds (1) does not simply distinguish between familiar and unfamiliar singers, (2) requires a small sample of both songs and song types, (3) does not rely on individual-specific sequences of song types and (4) is not likely to rely on group-specific vocal signatures potentially available in cooperatively breeding groups. We conclude that this is a case of true recognition and suggest that the lack of a dear enemy effect in this and other species with large repertoires may relate to the role of song in mate attraction and the perception of neighbours as a threat to future paternity.