Geographic song discrimination in relation to dispersal distances in song sparrows

Whether geographic variation in signals actually affects communication between individuals depends on whether discriminable differences in signals occur over distances that individuals move in their lifetimes. We measure the ability of song sparrows (Melospiza melodia) to discriminate foreign from local songs using foreign songs recorded at a series of increasing distances and compare the results with previous measurements of dispersal distances. We test discrimination in males using playback of songs on territories and measuring approach and in females using playback to estradiol-treated captives and measuring courtship display. Females fail to discriminate against foreign songs recorded at 18 km but do discriminate against foreign songs recorded at 34, 68, 135, and 540 km. Males fail to discriminate against foreign songs recorded at 18, 34, 68, 135, and 270 km but do discriminate against foreign songs from 540 km. Females are more discriminating, but even they do not discriminate at a distance three times the root-mean-square dispersal distance, as estimated from mtDNA variation. We suggest that female preference for local songs benefits females not because it allows them to reject foreign males but because accurate production of local song serves as a test of song-learning ability.     

Seasonal changes in song nuclei and song behavior in Gambel's white-crowned sparrows

In males of several songbird species, the morphology of forebrain nuclei that control song changes seasonally. The only seasonally breeding songbird in which seasonal changes in the structure of song control nuclei have been reported not to occur is the nonmigratory Nuttall's subspecies of white-crowned sparrow. In the present study, we manipulated photoperiod and plasma testosterone concentrations in captive male white-crowned sparrows of the migratory Gambel's subspeices. Males exposed to photoperiods and plasma testosterone concentrations typical of those experienced by wild breeding males had larger song control nuclei than males held on a winter photoperiod. We also found seasonal change in stereotypy of spectral and temporal parameters of song in wild Gambel's white-crowned sparrows. We hypothesize that seasonal changes in song control nuclei may correlate with seasonal changes in song stereotypy. © 1995 John Wiley & Sons, Inc.     

Androgens and estrogens induce seasonal-like growth of song nuclei in the adult songbird brain

In seasonally breeding songbirds, the brain regions that control song behavior undergo dramatic structural changes at the onset of each annual breeding season. As spring approaches and days get longer, gonadal testosterone (T) secretion increases and triggers the growth of several song control nuclei. T can be converted to androgenic and estrogenic metabolites by enzymes expressed in the brain. This opens the possibility that the effects of T may be mediated via the androgen receptor, the estrogen receptor, or both. To test this hypothesis, we examined the effects of two bioactive T metabolites on song nucleus growth and song behavior in adult male white-crowned sparrows. Castrated sparrows with regressed song control nuclei were implanted with silastic capsules containing either crystalline T, 5alpha-dihydrotestosterone (DHT), estradiol (E(2)), or a combination of DHT+E(2). Control animals received empty implants. Song production was highly variable within treatment groups. Only one of seven birds treated with E(2) alone was observed singing, whereas a majority of birds with T or DHT sang. After 37 days of exposure to sex steroids, we measured the volumes of the forebrain song nucleus HVc, the robust nucleus of the archistriatum (RA), and a basal ganglia homolog (area X). All three steroid treatments increased the volumes of these three song nuclei when compared to blank-implanted controls. These data demonstrate that androgen and estrogen receptor binding are sufficient to trigger seasonal song nucleus growth. These data also suggest that T's effects on seasonal song nucleus growth may depend, in part, upon enzymatic conversion of T to bioactive metabolites.     

Phonology and Geographic Song Discrimination in Song Sparrows

We investigated whether song sparrows discriminate foreign from local songs using specific phonologic markers, a mechanism of geographic discrimination previously described for some other songbirds. Song sparrows from Linesville, Pennsylvania (PA) respond more strongly to local songs than to songs from Millbrook, New York (NY). We identify 61 note categories in PA and NY songs, of which 13 are unique to PA, 17 are unique to NY, and 31 are shared. The most common note category in PA song, ‘buzz BO5’, is present in 89% of PA songs and only in 8% of NY songs; this difference in percentage representation is the largest we found for any note category. Substituting this potential PA marker into NY songs, however, did not make those songs more salient to PA sparrows; instead, PA males tested with territorial playback responded significantly less aggressively to NY/PA hybrid songs than to NY songs. A series of control experiments showed that song sparrows do not detect substitution of PA notes into PA songs or of NY notes into NY songs. The results weigh against the hypothesis that geographic discrimination in song sparrows is accomplished simply by recognition of a small number of phonologic markers.     

Species song discrimination in choice experiments with territorial male swamp and song sparrows

Playbacks of synthetic and normal songs were used to determine what song parameters are important to species song recognition by territorial male swamp sparrows (Melospiza georgiana) and song sparrows (Melospiza melodia). Syllable morphology influences the discrimination of conspecific song from song sparrow song in adult swamp sparrows, but temporal pattern does not; this result duplicates that found in studies of song learning of the young in this species. Species song recognition is influenced by both syllable morphology and temporal pattern in adult song sparrows, in contrast to the situation in young sparrows, which seem not to distinguish their own species' song from that of swamp sparrows on either cue.     

Androgens and estrogens induce seasonal‐like growth of song nuclei in the adult songbird brain

In seasonally breeding songbirds, the brain regions that control song behavior undergo dramatic structural changes at the onset of each annual breeding season. As spring approaches and days get longer, gonadal testosterone (T) secretion increases and triggers the growth of several song control nuclei. T can be converted to androgenic and estrogenic metabolites by enzymes expressed in the brain. This opens the possibility that the effects of T may be mediated via the androgen receptor, the estrogen receptor, or both. To test this hypothesis, we examined the effects of two bioactive T metabolites on song nucleus growth and song behavior in adult male white‐crowned sparrows. Castrated sparrows with regressed song control nuclei were implanted with silastic capsules containing either crystalline T, 5α‐dihydrotestosterone (DHT), estradiol (E₂), or a combination of DHT+E₂. Control animals received empty implants. Song production was highly variable within treatment groups. Only one of seven birds treated with E₂ alone was observed singing, whereas a majority of birds with T or DHT sang. After 37 days of exposure to sex steroids, we measured the volumes of the forebrain song nucleus HVc, the robust nucleus of the archistriatum (RA), and a basal ganglia homolog (area X). All three steroid treatments increased the volumes of these three song nuclei when compared to blank‐implanted controls. These data demonstrate that androgen and estrogen receptor binding are sufficient to trigger seasonal song nucleus growth. These data also suggest that T's effects on seasonal song nucleus growth may depend, in part, upon enzymatic conversion of T to bioactive metabolites. © 2003 Wiley Periodicals, Inc. J Neurobiol 57:130–140, 2003     

Territory establishment,song learning strategies and survival in song sparrows

In most songbirds, the processes of song learning and territory establishment overlap in the early life and a young bird usually winds up with songs matching those of his territorial neighbors in his first breeding season. In the present study, we examined the relationships among the timing of territory establishment, the pattern of song learning and territorial success in a sedentary population of song sparrows (Melospiza melodia). Males in this population tend to learn their songs from their neighbors and consequently they show high song sharing with neighbors and use these shared songs preferentially in interactions with them. Males also show significant variation in the timing of territory establishment, ranging from their natal summer to the next spring. Using a three-year dataset, we found that the timing of territory establishment did not systematically affect the composition of the song repertoire of the tutee: early establishers and late establishers learned equally as much from their primary tutors and had a similar number of tutors and similar repertoire sizes, nor did timing of territory establishment affect subsequent survival on territory. Therefore, the song-learning program of song sparrows seems versatile enough to lead to high song sharing even for birds that establish territories relatively late.     

Duration of melatonin regulates seasonal changes in song control nuclei of the house sparrow, Passer domesticus: independence from gonads and circadian entrainment

Avian behavior and physiology are temporally regulated by a complex circadian clock on both a daily and an annual basis. The circadian secretion of the hormone melatonin is a critical component of the regulation of circadian/daily processes in passerine birds, but there is little evidence that the gland regulates annual changes in primary reproductive function. Here it is shown that locomotor rhythms of house sparrows, Passer domesticus, which are made arrhythmic by either pinealectomy or maintenance in constant light, can be synchronized by daily administration of melatonin of different durations to simulate the melatonin profiles indicative of long and short photoperiods. Pinealectomized male sparrows maintained in constant darkness were entrained by both melatonin regimens. In both cases, testes were regressed and the song control nuclei were small. Intact male house sparrows maintained in constant light were also entrained to both melatonin regimens. However, sparrows that received a long duration melatonin cycle exhibited small song control nuclei, while sparrows that received short duration melatonin or no melatonin at all exhibited large song control nuclei. The data indicate that seasonal changes in melatonin duration contribute to the regulation of song control nuclei.     

Neighbour-stranger song discrimination in territorial ortolan bunting Emberiza hortulana males

Neighbour-stranger discrimination has been demonstrated in many species, but the mechanisms employed in discrimination vary. We tested whether an oscine bird with small repertoire size, the ortolan bunting Emberiza hortulana , discriminated between songs of neighbours and strangers. We performed playback experiments to measure response of males to a repeated single example of a single song type derived from a repertoire of a neighbour or stranger. Thirteen males were tested twice each, and in both cases songs were broadcast from the territory boundary shared by the subject male and the neighbour. Subjects responded more aggressively to songs of strangers than neighbours, i.e. they approached the loudspeaker faster and came closer and did more flights during the playback of stranger song. We found no significant differences in vocal response between treatments. We conclude that ortolan bunting can discriminate between songs of neighbours and strangers. This study provides experimental evidence for ortolan buntings in neighbour-stranger discrimination. It also demonstrates that a single example of song is enough to discriminate between neighbours and strangers. We discuss which song characteristics are the possible acoustic basis for discrimination in the studied species.     

Alternative forms of song matching in song sparrows

Song matching, replying to a song with a similar song, occurs in many songbird species. Almost all investigations of song matching have been of type matching, where one bird's reply is unambiguously similar to the other's song (i.e. the same song type). In many populations, however, neighbours do not share song types, and therefore cannot type-match. We hypothesized that a bird lacking a true type match could still song-match a stimulus song with a song from his repertoire that was similar in some way the birds recognized. We tested this hypothesis in song sparrows, Melospiza melodia, in two playback experiments. We played the subject a stranger song that was similar to one or more of his songs, but a type match to none of them. In the first experiment, we used playback songs that began with two buzzes (‘double-buzz’ songs). In the second experiment, we used songs that began with a slow trill that increased in tempo ('speed-up' songs). Birds replied at rates significantly above chance with their own double-buzz, or speed-up song match to the respective types of playback. The results suggest that birds who do not share true song types, can still song-match each other. This broad-sense form of song matching may also occur in populations with low song type sharing. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Testosterone treatment increases the metabolic capacity of adult avian song control nuclei

In songbirds, the size of brain nuclei that control song learning and production change seasonally. These changes are mainly controlled by seasonal changes in plasma testosterone (T) concentration. One hypothesis to explain why it may be adaptive for these areas to regress in the fall is that this would decrease the metabolic demand of maintaining a large song system when singing is reduced or absent. We used a marker for cellular metabolism to examine birds with regressed song nuclei and compared them to birds whose song nuclei were induced to grow by administration of exogenous T. Photorefractory male Gambel's white-crowned sparrows were captured during their autumnal migration and kept in outdoor aviaries on a natural photoperiod. We implanted birds with Silastic capsules containing T or with empty implants. Three weeks later the birds were sacrificed. We assayed the brains for cytochrome oxidase (CO) activity and measured the volume of four song nuclei: HVc, RA, 1MAN, and area X. All four nuclei increased in volume in response to T treatment. T treatment increased the metabolic capacity of area X, HVc, and RA relative to surrounding tissue but had no effect on the metabolic capacity of 1MAN. These results support the hypothesis that song nuclei are more metabolically active under the influence of T than they are when plasma T levels are low.     

Species song discrimination in adult female song and swamp sparrows

Six female song sparrows (Melospiza melodia) and six female swamp sparrows (Melospiza georgiana) were treated with oestradiol, and their response to song measured by frequency and intensity of copulation-solicitation display. Test stimuli were synthetic songs composed of either song sparrow or swamp sparrow syllables assembled in either song-sparrow-like or swamp-sparrow-like temporal patterns. Female song sparrows responded preferentially to songs containing their own species' syllables and to songs containing their own species' temporal patterns. Swamp sparrows were also sensitive to both syllable type and temporal, pattern, in contrast to male swamp sparrows, which show no preference for swamp sparrow temporal patterns.     

Neighbor-stranger discrimination by song in a suboscine bird, the alder flycatcher, Empidonax alnorum

Bird song and its functions have been studied extensively formore than 50 years, but almost entirely in oscine passerines.Few studies have investigated any aspect of song in suboscinepasserines. This is significant because song development andthe extent of individual variation in song differs greatly betweenthese groups. Learning and auditory feedback play major rolesin song development in all oscines studied, but apparently nopart in song ontogeny in suboscines. The ability of territorialoscine males to discriminate between songs of neighbors andstrangers has received considerable attention, but this phenomenonis virtually unstudied in suboscines. We tested whether a suboscinebird, the alder flycatcher (Empidonax alnorum), was able todiscriminate between songs of neighbors and strangers despitelimited individual variation in song. We performed playbackexperiments to measure responses of males to songs of neighborsand strangers broadcast from the territory boundary shared bythe subject and the neighbor. Subjects responded more aggressivelyto songs of strangers than to songs of neighbors. These resultsfurther our understanding of the evolution of song and its functionsin suboscines by demonstrating that, similar to their oscinerelatives, they can discriminate between the songs of neighborsand strangers.     

Age-related changes in song repertoire size and song type sharing in the Whinchat Saxicola rubetra

In many oscine passerines males’ songs, the repertoire size increases with age. At the same time it often remains unknown when and where males learn new songs. To infer the Whinchat Saxicola rubetra song learning strategy, we described and catalogued song-type repertoire, revealed age differences and examined song sharing strategies among neighbouring and distant males. We recorded song vocalizations of 40 males in a limited (104 ha) study plot during four years. Whinchats produce short and discrete songs with clear intersong pauses. In total 45 song types were allocated, the individual repertoire size averaged 23.5 ± 7.6 song types (range 9–34 song types). The males’ age significantly influenced the song-type repertoire size. The second calendar year (first breeding) males had a lower repertoire size than the older males. The majority of song types were shared by less than half of males in our sample. The Jaccard similarity indexes varied from 0.5 to 0.7. We could not find a relationship between males’ song sharing and geographic distances between their nests. We assume that Whinchat males learned new songs in the local population before territory establishment.     

The effect of auditory distractors on song discrimination in male canaries (Serinus canaria)

Male songbirds such as canaries produce complex learned vocalizations that are used in the context of mate attraction and territory defense. Successful mate attraction or territorial defense requires that a bird be able to recognize individuals based on their vocal performance and identify these songs in a noisy background. In order to learn more about how birds are able to solve this problem, we investigated, with a two-alternative choice procedure, the ability of adult male canaries to discriminate between conspecific song segments from two different birds and to maintain this discrimination when conspecific songs are superimposed with a variety of distractors. The results indicate that male canaries have the ability to discriminate, with a high level of accuracy song segments produced by two different conspecific birds. Song discrimination was partially maintained when the stimuli were masked by auditory distractors, but the accuracy of the discrimination progressively declined as a function of the number of masking distractors. The type of distractor used in the experiments (other conspecific songs or different types of artificial white noise) did not markedly affect the rate of deterioration of the song discrimination. These data indicate that adult male canaries have the perceptual abilities to discriminate and selectively attend to one ongoing sound that occurs simultaneously with one or more other sounds. The administration of a noradrenergic neurotoxin did not impair markedly the discrimination learning abilities although the number of subjects tested was too small to allow any firm conclusion. In these conditions, however, the noradrenergic lesion significantly increased the number failures to respond in the discrimination learning task suggesting a role, in canaries, of the noradrenergic system in some attentional processes underlying song learning and processing.     

Female canaries that respond and discriminate more between male songs of different quality have a larger song control nucleus (HVC) in the brain

In male songbirds the song control pathway in the forebrain is responsible for song production and learning. In most species, females do not sing and have smaller nuclei in the song control pathway. Although the function of the pathway in females is assumed to be associated with the perception of male song, there is little direct evidence to support this view. In this study on female canaries, we investigate the role of two key nuclei in the song control pathway (HVC and lMAN) in relation to playback of male song. Male canaries produce elaborate songs that function to attract and stimulate females. The songs are constructed from smaller units called syllables, and special syllables with a more complex structure (sexy syllables) are known to induce females to perform copulation solicitation displays (CSD) as an invitation to mate. By using computer-edited experimental songs, we first show that females discriminate between songs by producing significantly more CSD to those containing sexy syllables. We then sectioned the brains and used in situ hybridization to reveal song nuclei containing androgen receptors. We report positive correlations between the size of HVC and both total CSD response and the amount of discrimination between sexy and nonsexy songs. We found no such relationships between these measures and the size of lMAN. These results provide some evidence to support the view that, in female canaries HVC is involved in female perception and discrimination of male song. The results also have implications for the evolution of complex male songs by sexual selection and female choice.     

Social influences during song development in the song sparrow: a laboratory experiment simulating field conditions

Oscine songbirds are exposed to many more songs than they keep for their final song repertoire and little is known about how a bird selects the particular song(s) to sing as an adult. We simulated in the laboratory the key variables of the natural song learning environment and examined the song selection process in nine hand-reared male song sparrows, Melospiza melodia, a species in which males sing 5-11 song types. During their second and third months (their presumed sensitive period), subjects were rotated equally among four live adult male tutors that had been neighbours in the field. Tutors were housed in individual aviary 'territories' in four corners of the roof of a building; subjects could see only one tutor at a time, but they could hear the others at a short distance. Later in their first year (months 5-12), half the subjects were again rotated among all four tutors and the other half were randomly stationed next to just one tutor. Results from this experiment confirm and extend the findings from our two previous field studies of song learning in this species. Young males in this experiment (1) learned whole song types, (2) learned songs from multiple tutors, (3) preferentially learned songs that were shared among their tutors, (4) learned songs that other young males in their group also chose, and (5) learned more songs from the tutor they were stationed next to during the later stage (stationary subjects). These last two results support the late influence hypothesis that interactions after a bird's sensitive period affect song repertoire development. Copyright 2000 The Association for the Study of Animal Behaviour.     

Hearing in the swamp sparrow,Melospiza georgiana,and the song sparrow,Melospiza melodia

Operant conditioning techniques and a psychophysical tracking procedure were used to measure thresholds for pure tones in two congeneric species of sparrows: swamp sparrows and song sparrows. Thresholds were measured both in quiet conditions and in the presence of white noise. The two species were similar in their gross pattern of auditory sensitivity with most sensitive thresholds in the frequency region of 1·0–8·0 kHz and poor hearing above and below this range. However, the best sensitivity occurred at 2 kHz for the song sparrows and at 4 kHz for the swamp sparrows. This pattern was also observed in song spectra of the two species. The songs of song sparrows had more power in lower frequencies than the songs of swamp sparrows. Masked auditory thresholds for both species were very similar, showing a generally increasing signal-to-noise ratio with increasing frequency. The similarity between swamp and song sparrows in these measures of basic auditory sensitivity suggests that the primary mechanisms underlying selective vocal learning in these sparrows probably reside at higher levels than the peripheral auditory sensitivities.     

Male response to historical and geographical variation in bird song

In many species, individuals discriminate among sexual signals of conspecific populations in the contexts of mate choice and male–male competition. Differences in signals among populations (geographical variation) are in part the result of signal evolution within populations (temporal variation). Understanding the relative effect of temporal and geographical signal variation on signal salience may therefore provide insight into the evolution of behavioural discrimination. However, no study, to my knowledge, has compared behavioural response to historical signals with response to current signal variation among populations. Here, I measured the response of male white-crowned sparrows (Zonotrichia leucophrys) to historical songs compared with current songs from their local population, a nearby non-local population and a distant population. Males responded most strongly to current local songs, less, but equally, to historical local and current non-local songs, and least to songs of the distant population. Moreover, response to both temporal and geographical variation in song was proportional to how much songs differed acoustically from current local songs. Signal evolution on an ecological time scale appears to have an effect on signal salience comparable to differences found between current neighbouring populations, supporting the idea that behavioural discrimination among learned signals of conspecific populations can evolve relatively rapidly.     

Typical versions of learned swamp sparrow song types are more effective signals than are less typical versions

The learned songs of songbirds often cluster into population-wide types. Here, we test the hypothesis that male and female receivers respond differently to songs depending on how typical of those types they are. We used computational methods to cluster a large sample of swamp sparrow (Melospiza georgiana) songs into types and to estimate the degree to which individual song exemplars are typical of these types. We then played exemplars to male and female receivers. Territorial males responded more aggressively and captive females performed more sexual displays in response to songs that are highly typical than to songs that are less typical. Previous studies have demonstrated that songbirds distinguish song types that are typical for their species, or for their population, from those that are not. Our results show that swamp sparrows also discriminate typical from less typical exemplars within learned song-type categories. In addition, our results suggest that more typical versions of song types function better, at least in male–female communication. This finding is consistent with the hypothesis that syllable type typicality serves as a proxy for the assessment of song learning accuracy.