Eding [6]. In many species males modulate their song in an aggressive context: they might select certain song types matching a rival [7], or produce specific song elements only in situations of high arousal [8]. In addition, birds can change song characteristics such as frequency patterns and trill 1326631 rate [9,10]. Male as well as female listeners respond differentiated to such modulations [11?4]. Song modulations can occur on two domains: on the one hand, birds may change the general output of song (e.g. song rate oramplitude), i.e. measures that potentially every male can vary within broad limits. On the other hand, modulation also occurs in structural song characteristics. Structural characteristics describe, for example, song repertoire characteristics [15] or song parts that are challenging to sing, such as rapid broadband trills (reviewed in [16]), specific song trills [17] or consistent NT 157 site syllables [18]. Structural song patterns have been classified as `index signals’ that honestly communicate a physical trait related to male quality [19]. Only very few studies have revealed a capability of individuals to modulate such physically constrained signals within narrow limits [9,10,20,21]. Thus, from a functional point of view, index signals such as structural song parameters should play an important role in the communication of competitive ability. The steroid hormone LED 209 testosterone plays an important role in the regulation of adult singing and territorial behaviors and the associated vocalizations during breeding are facilitated by testosterone in a wide range of male vertebrates (reviewed in [22], [23]). Therefore, it has been suggested that testosterone might play an important role in resource allocation for competitive behaviorTestosterone Affects Song Modulationduring reproduction (reviewed in [24]). From this point of view, testosterone should act specifically on signals that communicate the motivation or ability of individuals to engage in competitive situations and is, therefore, expected to be involved in contextdependent adjustment of such signals. However, details of the interplay between hormones, territorial aggression and signal plasticity in a natural context are largely unknown. Manipulations of testosterone levels may alter song output (measured, for example, as song rate or duration; e.g. [25?9]). Whether testosterone also affects structural song parameters is less clear. In barn swallows (Hirundo rustica), the duration and pulse rate of the harsh `rattle’ element correlated moderately with absolute testosterone levels [30]. Manipulation studies suggested that zebra finches (Taeniopygia guttata) treated with testosterone decreased the fundamental frequency of harmonic stacks in their song [31]. Other correlational and experimental studies with testosterone treatment failed to find effects on structural song parameters [29,32,33]. Studies that implant birds with testosterone may be problematic, because especially immediately after implantation testosterone may circulate in pharmacological doses [34,35]. It is thus questionable whether manipulations exclusively within the physiological range of testosterone would reveal similar results. Treatments inhibiting the action of testosterone or its major metabolite estradiol by blocking the androgen receptor and/or the conversion to estradiol avoid such pharmacological effects (but can only inhibit, not enhance effects of steroid hormones). The ?so far – only study in which the andro.Eding [6]. In many species males modulate their song in an aggressive context: they might select certain song types matching a rival [7], or produce specific song elements only in situations of high arousal [8]. In addition, birds can change song characteristics such as frequency patterns and trill 1326631 rate [9,10]. Male as well as female listeners respond differentiated to such modulations [11?4]. Song modulations can occur on two domains: on the one hand, birds may change the general output of song (e.g. song rate oramplitude), i.e. measures that potentially every male can vary within broad limits. On the other hand, modulation also occurs in structural song characteristics. Structural characteristics describe, for example, song repertoire characteristics [15] or song parts that are challenging to sing, such as rapid broadband trills (reviewed in [16]), specific song trills [17] or consistent syllables [18]. Structural song patterns have been classified as `index signals’ that honestly communicate a physical trait related to male quality [19]. Only very few studies have revealed a capability of individuals to modulate such physically constrained signals within narrow limits [9,10,20,21]. Thus, from a functional point of view, index signals such as structural song parameters should play an important role in the communication of competitive ability. The steroid hormone testosterone plays an important role in the regulation of adult singing and territorial behaviors and the associated vocalizations during breeding are facilitated by testosterone in a wide range of male vertebrates (reviewed in [22], [23]). Therefore, it has been suggested that testosterone might play an important role in resource allocation for competitive behaviorTestosterone Affects Song Modulationduring reproduction (reviewed in [24]). From this point of view, testosterone should act specifically on signals that communicate the motivation or ability of individuals to engage in competitive situations and is, therefore, expected to be involved in contextdependent adjustment of such signals. However, details of the interplay between hormones, territorial aggression and signal plasticity in a natural context are largely unknown. Manipulations of testosterone levels may alter song output (measured, for example, as song rate or duration; e.g. [25?9]). Whether testosterone also affects structural song parameters is less clear. In barn swallows (Hirundo rustica), the duration and pulse rate of the harsh `rattle’ element correlated moderately with absolute testosterone levels [30]. Manipulation studies suggested that zebra finches (Taeniopygia guttata) treated with testosterone decreased the fundamental frequency of harmonic stacks in their song [31]. Other correlational and experimental studies with testosterone treatment failed to find effects on structural song parameters [29,32,33]. Studies that implant birds with testosterone may be problematic, because especially immediately after implantation testosterone may circulate in pharmacological doses [34,35]. It is thus questionable whether manipulations exclusively within the physiological range of testosterone would reveal similar results. Treatments inhibiting the action of testosterone or its major metabolite estradiol by blocking the androgen receptor and/or the conversion to estradiol avoid such pharmacological effects (but can only inhibit, not enhance effects of steroid hormones). The ?so far – only study in which the andro.
