Sarah Kiefer

Associated and disassociated patterns in hormones, song, behavior and brain receptor expression between life-cycle stages in male black redstarts, Phoenicurus ochruros

Testosterone has been suggested to be involved in the regulation of male territorial behavior. For example, seasonal peaks in testosterone typically coincide with periods of intense competition between males for territories and mating partners. However, some species also express territorial behavior outside a breeding context when testosterone levels are low and, thus, the degree to which testosterone facilitates territorial behavior in these species is not well understood. We studied territorial behavior and its neuroendocrine correlates in male black redstarts. Black redstarts defend territories in spring during the breeding period, but also in the fall outside a reproductive context when testosterone levels are low. In the present study we assessed if song output and structure remain stable across life-cycle stages. Furthermore, we assessed if brain anatomy may give insight into the role of testosterone in the regulation of territorial behavior in black redstarts. We found that males sang spontaneously at a high rate during the nonbreeding period when testosterone levels were low; however the trill-like components of spontaneously produced song contained less repetitive elements during nonbreeding than during breeding. This higher number of repetitive elements in trills did not, however, correlate with a larger song control nucleus HVC during breeding. However, males expressed more aromatase mRNA in the preoptic area - a brain nucleus important for sexual and aggressive behavior - during breeding than during nonbreeding. In combination with our previous studies on black redstarts our results suggest that territorial behavior in this species only partly depends on sex steroids: spontaneous song output, seasonal variation in trills and non-vocal territorial behavior in response to a simulated territorial intruder seem to be independent of sex steroids. However, context-dependent song during breeding may be facilitated by testosterone - potentially by conversion of testosterone to estradiol in the preoptic area.

Buzzwords in Females’ Ears? The Use of Buzz Songs in the Communication of Nightingales (Luscinia megarhynchos)

Differences in individual male birds’ singing may serve as honest indicators of male quality in male-male competition and female mate choice. This has been shown e.g. for overall song output and repertoire size in many bird species. More recently, differences in structural song characteristics such as the performance of physically challenging song components were analysed in this regard. Here we show that buzz elements in the song of nightingales (Luscinia megarhynchos) hold the potential to serve as indicators of male quality and may therefore serve a communicative function. Buzzes were produced with considerable differences between males. The body weight of the males was correlated with one measure of these buzzes, namely the repetition rate of the buzz subunits, and individuals with larger repertoires sang buzzes at higher subunit-rates. A model of buzz performance constraints suggested that buzzes were sung with different proficiencies. In playback experiments, female nightingales showed more active behaviour when hearing buzz songs. The results support the idea that performance differences in the acoustic fine structure of song components are used in the communication of a large repertoire species such as the nightingale.

Testosterone Affects Song Modulation during Simulated Territorial Intrusions in Male Black Redstarts (Phoenicurus ochruros)

Although it has been suggested that testosterone plays an important role in resource allocation for competitive behavior, details of the interplay between testosterone, territorial aggression and signal plasticity are largely unknown. Therefore, we investigated if testosterone acts specifically on signals that communicate the motivation or ability of individuals to engage in competitive situations in a natural context. We studied the black redstart, a territorial songbird species, during two different life-cycle stages, the early breeding phase in spring and the non-breeding phase in fall. Male territory holders were implanted with the androgen receptor blocker flutamide (Flut) and the aromatase inhibitor letrozole (Let) to inhibit the action of testosterone and its estrogenic metabolites. Controls received a placebo treatment. Three days after implantation birds were challenged with a simulated territorial intrusion (STI). Song was recorded before, during and after the challenge. In spring, both treatment groups increased the number of elements sung in parts of their song in response to the STI. However, Flut/Let-implanted males reacted to the STI with a decreased maximum acoustic frequency of one song part, while placebo-implanted males did not. Instead, placebo-implanted males sang the atonal part of their song with a broader frequency range. Furthermore, placebo-, but not Flut/Let-implanted males, sang shorter songs with shorter pauses between parts in the STIs. During simulated intrusions in fall, when testosterone levels are naturally low in this species, males of both treatment groups sang similar to Flut/Let-implanted males during breeding. The results suggest that song sung during a territorial encounter is of higher competitive value than song sung in an undisturbed situation and may, therefore, convey information about the motivation or quality of the territory holder. We conclude that testosterone facilitates context-dependent changes in song structures that may be honest signals of male quality in black redstarts.

Life-history and hormonal control of aggression in black redstarts: Blocking testosterone does not decrease territorial aggression, but changes the emphasis of vocal behaviours during simulated territorial intrusions

IntroductionMany studies in behavioural endocrinology attempt to link territorial aggression with testosterone, but the exact relationship between testosterone and territorial behaviour is still unclear and may depend on the ecology of a species. The degree to which testosterone facilitates territorial behaviour is particularly little understood in species that defend territories during breeding and outside the breeding season, when plasma levels of testosterone are low. Here we suggest that species that defend territories in contexts other than reproduction may have lost the direct regulation of territorial behaviour by androgens even during the breeding season. In such species, only those components of breeding territoriality that function simultaneously as sexually selected signals may be under control of sex steroids.ResultsWe investigated black redstarts (Phoenicurus ochruros), a species that shows periods of territoriality within and outside of the breeding season. We treated territorial males with an anti-androgen and an aromatase inhibitor during the breeding season to block both the direct and indirect effects of testosterone. Three and ten days after the treatment, implanted males were challenged with a simulated territorial intrusion. The treatment did not reduce the overall territorial response, but it changed the emphasis of territoriality: experimental males invested more in behaviours addressed directly towards the intruder, whereas placebo-treated males put most effort into their vocal response, a component of territoriality that may be primarily directed towards their mating partner rather than the male opponent.ConclusionsIn combination with previous findings, these data suggest that overall territoriality may be decoupled from testosterone in male black redstarts. However, high levels of testosterone during breeding may facilitate-context dependent changes in song.

Does age matter in song bird vocal interactions? Results from interactive playback experiments

The song of oscines provides an extensively studied model of age-dependent behaviour changes. Male and female receivers might use song characteristics to obtain information about the age of a signaller, which is often related to its quality. Whereas most of the age-dependent song changes have been studied in solo singing, the role of age in vocal interactions is less well understood. We addressed this issue in a playback study with common nightingales (Luscinia megarhynchos). Previous studies showed that male nightingales had smaller repertoires in their first year than older males and males adjusted their repertoire towards the most common songs in the breeding population. We now compared vocal interaction patterns in a playback study in 12 one year old and 12 older nightingales (cross-sectional approach). Five of these males were tested both in their first and second breeding season (longitudinal approach). Song duration and latency to respond did not differ between males of different ages in either approach. In the cross-sectional approach, one year old nightingales matched song types twice as often as did older birds. Similarly, in the longitudinal approach all except one bird reduced the number of song type matches in their second season. Individuals tended to overlap songs at higher rates in their second breeding season than in their first. The higher levels of song type matches in the first year and song overlapping by birds in their second year suggest that these are communicative strategies to establish relationships with competing males and/or choosy females.

Learn it now, sing it later? Field and laboratory studies on song repertoire acquisition and song use in nightingales

In many bird species, song changes with age. The mechanisms that account for such changes are only partially understood. Common nightingales Luscinia megarhynchos change the size and composition of their repertoire between their first and second breeding season. To inquire into mechanisms involved in such changes, we compared the singing of 1-year-old and older free-living nightingales. Older males have more song types in common than have 1-year olds. Certain song types frequently sung by older birds did not (or only rarely) occur in the repertoire of yearlings (‘mature’ song types). We conducted learning experiments with hand-reared nightingales to address reasons for the lack of mature song types. The acquisition success of mature songs was as good as that of control songs (commonly sung by both age groups). However, the analysis of song type use revealed that all yearlings sang common song types more often than mature types. This indicates that the absence of certain song types in the repertoires of free-living yearlings cannot be accounted for by learning and/or motor constraints during song learning. Moreover, our results suggest that in communication networks, animals may restrict the actual use of their signal repertoire to a certain subset depending on the context.

Female calling? Song responses to conspecific call playbacks in nightingales, Luscinia megarhynchos

A crucial issue for understanding the evolution, functions and mechanisms of complex communicative signals such as birdsong is to disentangle signal structures that serve to convey information in different contexts. The two main singing contexts of European male songbirds are the attraction of a female and the defence of a territory from other males. The method of choice to experimentally investigate the use of song in male–male interactions is to conduct playback studies, in which song is played to a resident male. Responses to the presence of a female, however, are hardly testable this way, since females of most European bird species do not sing. However, females do call in several situations. In a playback experiment, we investigated whether free-ranging male nightingales change their nocturnal singing in response to playbacks consisting of calls produced by either female or male conspecifics. In both cases, nightingales changed their singing style as compared to their singing before playback, and these changes were different in response to male and female calls. Males sang fewer whistle songs after male call playbacks and started to produce ‘initial whistles’ earlier when hearing female calls. Male call playbacks also led to an increase in song duration. An analysis of the call characteristics of both sexes used in the playbacks uncovered acoustic parameters that may account for the differentiated response. We conclude that male nightingales distinguish between female and male calls, and discuss the song characteristics that might be used to specifically address either males or females. Call playbacks proved to be useful for disentangling song characteristics that serve the multiple functions of birdsong.