Nicole Geberzahn

Bird song and anthropogenic noise: vocal constraints may explain why birds sing higher-frequency songs in cities

When animals live in cities, they have to adjust their behaviour and life histories to novel environments. Noise pollution puts a severe constraint on vocal communication by interfering with the detection of acoustic signals. Recent studies show that city birds sing higher-frequency songs than their conspecifics in non-urban habitats. This has been interpreted as an adaptation to counteract masking by traffic noise. However, this notion is debated, for the observed frequency shifts seem to be less efficient at mitigating noise than singing louder, and it has been suggested that city birds might use particularly high-frequency song elements because they can be produced at higher amplitudes. Here, we present the first phonetogram for a songbird, which shows that frequency and amplitude are strongly positively correlated in the common blackbird (Turdus merula), a successful urban colonizer. Moreover, city blackbirds preferentially sang higher-frequency elements that can be produced at higher intensities and, at the same time, happen to be less masked in low-frequency traffic noise.

Honey bees transfer olfactory memories established during flower visits to a proboscis extension paradigm in the laboratory

To see what effects learning in one context can exercise upon subsequent learning in other contexts the relation between olfactory learning performance of honey bees,Apis melliferaL., in a restrained classical conditioning paradigm for the proboscis extension reflex and chemosensory experience under natural circumstances was investigated. The effects of food-source odours and of the hives odour on subsequent laboratory proboscis extension performance were tested. There was a transfer effect for odours from a food source to proboscis extension reflex conditioning in the laboratory: bees extended their proboscis in response to food-source odour and showed resistance to its extinction. Hive odour had no detectable effect. Thus, the observed transfer effect for food-source odours was likely to be caused by memories established during foraging rather than by within-hive experience with the scent of harvested food. The observed transfer effect for odours from a food source thus points to a surprisingly weak role of context for the retrieval of olfactory memories established during foraging. Furthermore, it offers an experience-based explanation for variability in honey bee proboscis extension learning.

Females alter their song when challenged in a sex-role reversed bird species

Birdsong serves to attract mates and to deter territorial rivals. Even though song is not restricted to males, this dual function has almost exclusively been demonstrated for male song. To test the generality of hypotheses on birdsong, we investigated female song in the sex-role reversed, classically polyandrous African black coucal (Centropus grillii) in the context of female–female competition. We compared spontaneously vocalizing females with females vocally responding to a playback simulating a conspecific intruder. Females changed vocal parameters in response to playbacks: They lowered the pitch of their vocalizations and enhanced the duration of song elements when being challenged. Also, the composition of the vocalizations was altered. There was no significant correlation between pitch and body size parameters in spontaneous song, but there was for response songs, with larger females having a lower pitch. These changes in vocal properties suggest that the vocalizations are important for mutual assessment of competitive abilities in females. Our findings confirm the general role of intrasexual competition in vocal communication of birds.

Latent song type memories are accessible through auditory stimulation in a hand-reared songbird

The production of learned vocalizations such as in birdsong is often used to judge whether stimuli had been memorized upon their presentation. However, failures in the imitation of certain song patterns may also reflect impaired development of motor programmes or impaired memory retrieval rather than failures in stimulus memorization during auditory acquisition. To study this issue, we confronted adult hand-reared nightingales, Luscinia megarhynchos, with interactive playback experiments and used vocal matching as a behavioural tool to investigate their song type memories. Vocal matching is a common pattern-specific response that songbirds use in territorial countersinging. We distinguished two forms of pattern-specific matching: (1) song type matching (i.e. a bird replied with the same song type as the stimulus song), and (2) song group matching (i.e. the bird replied with a different song type which was, however, sequentially associated with the playback song presented earlier, i.e. during the tutoring). Some subjects used both song type and song group matching in response to song types they had not imitated from the tutor programme prior to the playback experiments. Our results indicate that nightingales store more song types in their sensory phase than they spontaneously recall from memory as adults. That is, memories of song types that were not performed in overt behaviour could be activated by vocal interactions, here induced by the interactive playback.

How a songbird with a continuous singing style modulates its song when territorially challenged

In songbirds of the temperate zone, often only males sing and their songs serve to attract females and to deter territorial rivals. In many species, males vary certain aspects of their singing behavior when engaged in territorial interactions. Such variation may be an honest signal of the traits of the signaler, such as fighting strength, condition, or aggressive motivation, and may be used by receivers in decisions on whether to retreat or to escalate a fight. This has been studied intensively in species that sing discontinuously, in which songs are alternating with silent pauses. We studied contextual variation in the song of skylarks (Alauda arvensis), a songbird with a large vocal repertoire and a continuous and versatile singing style. We exposed subjects to simulated territorial intrusions by broadcasting conspecific song and recorded their vocal responses. We found that males sing differently if they are singing spontaneously with no other conspecific around than if they are territorially challenged. In this last case, males produced lower-frequency syllables. Furthermore, they increased the sound density of their song: they increased the proportion of sound within song. They seem to do so by singing different elements of their repertoire when singing reactively. Furthermore, they increased the consistency of mean peak frequency: they repeated syllable types with less variability when singing reactively. Such contextual variation suggests that skylarks might use low frequencies, sound density, and song consistency to indicate their competitive potential, and thus, those song features might be important for mutual assessment of competitive abilities.

Assessing vocal performance in complex birdsong: a novel approach

BackgroundVocal performance refers to the ability to produce vocal signals close to physical limits. Such motor skills can be used by conspecifics to assess a signaler’s competitive potential. For example it is difficult for birds to produce repeated syllables both rapidly and with a broad frequency bandwidth. Deviation from an upper-bound regression of frequency bandwidth on trill rate has been widely used to assess vocal performance. This approach is, however, only applicable to simple trilled songs, and even then may be affected by differences in syllable complexity.ResultsUsing skylarks (Alauda arvensis) as a birdsong model with a very complex song structure, we detected another performance trade-off: minimum gap duration between syllables was longer when the frequency ratio between the end of one syllable and the start of the next syllable (inter-syllable frequency shift) was large. This allowed us to apply a novel measure of vocal performance - vocal gap deviation: the deviation from a lower-bound regression of gap duration on inter-syllable frequency shift. We show that skylarks increase vocal performance in an aggressive context suggesting that this trait might serve as a signal for competitive potential.ConclusionsWe suggest using vocal gap deviation in future studies to assess vocal performance in songbird species with complex structure.

Undirected (Solitary) Birdsong in Female and Male Blue-Capped Cordon-Bleus (Uraeginthus cyanocephalus) and Its Endocrine Correlates

Background Birdsong is a popular model system in research areas such as vocal communication, neuroethology or neuroendocrinology of behaviour. As most research has been conducted on species with male-only song production, the hormone-dependency of male song is well established. However, female singing and its mechanisms are poorly understood. Methodology/Principal Findings We characterised the song and its endocrine correlates of blue-capped cordon-bleus (Uraeginthus cyanocephalus), a species in which both sexes sing. Like other estrildids, they produce directed song during courtship and undirected (or solitary) song in isolation, i.e. when the mate is not visible or absent. We compare solitary song of blue-capped cordon-bleus to published descriptions of the song of its relative, the zebra finch (Taeniopygia guttata). Solitary song of cordon-bleus shared some overall song features with that of zebra finches but differed in spectro-temporal song features, sequential stereotypy and sequential organisation. The song of cordon-bleus was dimorphic with respect to the larger size of syllable repertoires, the higher song duration and the lower variability of pitch goodness (measuring the pureness of harmonic sounds) in males. However, in both sexes the overall plasma testosterone concentrations were low (ca. 300 pg/ml) and did not correlate with the sexually dimorphic song motor pattern. Despite such low concentrations, the increase in the rate of solitary song coincided with an increase in the level of testosterone. Furthermore, the latency to start singing after the separation from the mate was related to hormone levels. Conclusions/Significance Our findings suggest that the occurrence of solitary song but not its motor pattern might be under the control of testosterone in female and male cordon-bleus.

Rules of song development and their use in vocal interactions by birds with large repertoires.

Songbirds are well known for settling their disputes by vocal signals, and their singing plays a dominant role. Most studies on this issue have concentrated on bird species that develop and use small vocal repertoires. In this article we will go farther and focus on examples of how species with large song repertoires make use of their vocal competence. In particular, we will outline the study of interaction rules which have been elucidated by examining time- and pattern-specific relationships between signals exchanged by territorial neighbors. First we present an inquiry into the rules of song learning and development. In birds with large song repertoires, the ontogeny of such rules proceeds along a number of trajectories which help in understanding the often remarkable accomplishments of adult birds. In both approaches, our model species will be the Common Nightingale Luscinia megarhynchos that has been investigated intensively in the field and in the laboratory.

Mismatch in sexual dimorphism of developing song and song control system in blue-capped cordon-bleus, a songbird species with singing females and males

Brain song control regions of adult passerine birds are sexually dimorphic in species such as the zebra finch (Taeniopygia guttata) in which males sing whereas females do not. In many tropical bird species, however, females sing as well. Here we study for the first time the ontogeny of the song control system and the song in a species, in which both male and female sing regularly. In blue-capped cordon-bleus (Uraeginthus cyanocephalus), a distant relative of the zebra finch, both males and females start singing at around 30-40 day post-hatching (dph). First we quantified that sex-specific differences in song features emerged only in adulthood, after 250 dph of age: Adult females sang complex songs, which were slightly shorter and contained fewer syllables as compared to the males. Second, the development of forebrain song control regions HVC (proper name) and RA (nucleus robustus arcopallii) of blue-capped cordon-bleus was quantified in both sexes at 20, 30, 50, 100, 150, 250 dph as well as in old adults. The volume and neuron numbers of the HVC and RA were sexually dimorphic throughout the entire development and remained sexually dimorphic in adulthood. Since singing developed in a non sex-specific way until 250 dph, neural sex differences to a large extend precede the behavioral (song) sex differences. This suggests that these neuroanatomical sex differences are not causally related to the sexual differentiation of song patterns in this species.

Effects of behavioural time budgets and nest-building efficiency on male reproductive performance in red bishops ( Euplectes orix )

In this study we have investigated the effect of nest-building behaviour, courtship behaviour, and male–male interactions on male reproductive performance of the red bishop (Euplectes orix), a highly polygynous, colonially breeding weaverbird species. Previous studies on red bishops have revealed that male reproductive success is mainly determined by the number of nests built in a territory, and that reproductive success and the number of nests both vary substantially among males. Here we test whether the high variation observed in reproductive performance is related to male behavioural patterns (as assessed by time-budget analysis) and/or nest-building efficiency (as assessed by the number of fibres incorporated in the nest in unit time). We collected data on male time budgets and analysed behavioural patterns for three breeding seasons. A greater number of nests built was positively related to both the amount of time allocated to nest-building behaviour and the efficiency of nest building. Neither the amount of time spent in courtship behaviour nor the amount of time spent in male–male interactions was related to reproductive success. Male reproductive success, irrespective of the number of nests built, was partly determined by the number of fibres incorporated, suggesting some importance of nest quality in terms of nest chamber density to male reproductive success. There were no trade-offs with regard to time allocated to different behaviour. Instead, males differed with regard to their territory attendance and activity levels, which might be because of differences in energy resources and may thus reflect inherent differences in male body condition and, ultimately, male genetic quality.