C frequency of one song part in response to the STI, while placebo-implanted males kept this acoustic measure constant throughout the challenge. Furthermore, placebo-implanted males sang the atonal part of their song with a broader frequency range. In contrast to Flut/Let males, placebo-implanted males increased signal density by singing shorter songs with shorter pauses Title Loaded From File between song parts in the STI. In summary, these results provide a good example of the activational role of testosterone not only on song activity in general, but also on the specific singing style depending on the context. The results of this study indicate 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. 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. We conclude that these changes in song in response to a simulated territorial intruder were influenced by the Flut/Let treatment and by season: structural changes in song were less pronounced in Flut/Let males and in all males during non-breeding in fall compared to placebo-implanted males in spring.Song Modulation during Territorial ChallengesBlack redstarts of both treatment groups in spring sang more elements in parts A and C and placebo-implanted birds increased 1379592 the frequency Title Loaded From File bandwidth of part B when a simulated rival intruded the territory. Additionally, Flut/Let males decreased the maximum frequency of part A. These structural song parameters have been suggested to be physically challenging in other species (reviewed in [16]). Also, with regard to trilled parts, it has been suggested previously that the production of repeated (trilled) syllables with a high frequency bandwidth is challenging (reviewed in [16]). For example, in swamp sparrows, male age, size, and early developmental conditions correlated with these song characteristics, and can therefore serve as honest signals of male quality [14,56,5]. Females of some species prefer songs sung with a high trill rate and broad frequency bandwidth [12,57]. Furthermore, swamp sparrows increase both trill rate and frequency bandwidth in response to simulated territorial intruders [10]. Even though songs of control males were shorter during the STI than before (which might occur counter-intuitive at first, since usually birds increase song output when challenged), this resulted in a higher signal density. Increasing the signal density by changing the song output in an aggressive context seems to be a common strategy among bird species (e.g. [30,58]). In our study on black redstarts, this increase was realized by a shortening of pauses between song parts.Song in FallIn both treatment groups focal males sang fewer songs during and after the STI than before the experimental challenge (Fig. 2b, Table 2). Males of both treatment groups increased the number of elements in part A (Fig. 4c) and C (Fig. 4d) in response to the experimental challenge while decreasing the maximum frequency of part A (Fig. 4a) and decreasing the frequency bandwidth of part C (Table 2). Males sang part B with a significantly higher maximum frequency in response to the simulated territorial intrusion than during spontaneous song and this did again not significantly differ between placebo and Flut/Let-.C frequency of one song part in response to the STI, while placebo-implanted males kept this acoustic measure constant throughout the challenge. Furthermore, placebo-implanted males sang the atonal part of their song with a broader frequency range. In contrast to Flut/Let males, placebo-implanted males increased signal density by singing shorter songs with shorter pauses between song parts in the STI. In summary, these results provide a good example of the activational role of testosterone not only on song activity in general, but also on the specific singing style depending on the context. The results of this study indicate 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. 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. We conclude that these changes in song in response to a simulated territorial intruder were influenced by the Flut/Let treatment and by season: structural changes in song were less pronounced in Flut/Let males and in all males during non-breeding in fall compared to placebo-implanted males in spring.Song Modulation during Territorial ChallengesBlack redstarts of both treatment groups in spring sang more elements in parts A and C and placebo-implanted birds increased 1379592 the frequency bandwidth of part B when a simulated rival intruded the territory. Additionally, Flut/Let males decreased the maximum frequency of part A. These structural song parameters have been suggested to be physically challenging in other species (reviewed in [16]). Also, with regard to trilled parts, it has been suggested previously that the production of repeated (trilled) syllables with a high frequency bandwidth is challenging (reviewed in [16]). For example, in swamp sparrows, male age, size, and early developmental conditions correlated with these song characteristics, and can therefore serve as honest signals of male quality [14,56,5]. Females of some species prefer songs sung with a high trill rate and broad frequency bandwidth [12,57]. Furthermore, swamp sparrows increase both trill rate and frequency bandwidth in response to simulated territorial intruders [10]. Even though songs of control males were shorter during the STI than before (which might occur counter-intuitive at first, since usually birds increase song output when challenged), this resulted in a higher signal density. Increasing the signal density by changing the song output in an aggressive context seems to be a common strategy among bird species (e.g. [30,58]). In our study on black redstarts, this increase was realized by a shortening of pauses between song parts.Song in FallIn both treatment groups focal males sang fewer songs during and after the STI than before the experimental challenge (Fig. 2b, Table 2). Males of both treatment groups increased the number of elements in part A (Fig. 4c) and C (Fig. 4d) in response to the experimental challenge while decreasing the maximum frequency of part A (Fig. 4a) and decreasing the frequency bandwidth of part C (Table 2). Males sang part B with a significantly higher maximum frequency in response to the simulated territorial intrusion than during spontaneous song and this did again not significantly differ between placebo and Flut/Let-.
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