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1.  Song environment affects singing effort and vasotocin immunoreactivity in the forebrain of male Lincoln’s sparrows 
Hormones and behavior  2010;58(3):544-553.
Male songbirds often establish territories and attract mates by singing, and some song features can reflect the singer’s condition or quality. The quality of the song environment can change, so male songbirds should benefit from assessing the competitiveness of the song environment and appropriately adjusting their own singing behavior and the neural substrates by which song is controlled. In a wide range of taxa social modulation of behavior is partly mediated by the arginine vasopressin or vasotocin (AVP/AVT) systems. To examine the modulation of singing behavior in response to the quality of the song environment we compared the song output of laboratory-housed male Lincoln’s sparrows (Melospiza lincolnii) exposed to one week of chronic playback of songs categorized as either high or low quality, based on song length, complexity and trill performance. To explore the neural basis of any facultative shifts in behavior, we also quantified the subjects’ AVT immunoreactivity (AVT-IR) in three forebrain regions that regulate socio-sexual behavior: the medial bed nucleus of the stria terminalis (BSTm), the lateral septum (LS) and the preoptic area. We found that high quality songs increased singing effort and reduced AVT-IR in the BSTm and LS, relative to low quality songs. The effect of the quality of the song environment on both singing effort and forebrain AVT-IR raises the hypothesis that AVT within these brain regions plays a role in the modulation of behavior in response to competition that individual males may assess from the prevailing song environment.
PMCID: PMC2916047  PMID: 20399213
arginine vasotocin (AVT); bird song; competition; sexual selection; social behavior network
2.  Social Context–Induced Song Variation Affects Female Behavior and Gene Expression 
PLoS Biology  2008;6(3):e62.
Social cues modulate the performance of communicative behaviors in a range of species, including humans, and such changes can make the communication signal more salient. In songbirds, males use song to attract females, and song organization can differ depending on the audience to which a male sings. For example, male zebra finches (Taeniopygia guttata) change their songs in subtle ways when singing to a female (directed song) compared with when they sing in isolation (undirected song), and some of these changes depend on altered neural activity from a specialized forebrain-basal ganglia circuit, the anterior forebrain pathway (AFP). In particular, variable activity in the AFP during undirected song is thought to actively enable syllable variability, whereas the lower and less-variable AFP firing during directed singing is associated with more stereotyped song. Consequently, directed song has been suggested to reflect a “performance” state, and undirected song a form of vocal motor “exploration.” However, this hypothesis predicts that directed–undirected song differences, despite their subtlety, should matter to female zebra finches, which is a question that has not been investigated. We tested female preferences for this natural variation in song in a behavioral approach assay, and we found that both mated and socially naive females could discriminate between directed and undirected song—and strongly preferred directed song. These preferences, which appeared to reflect attention especially to aspects of song variability controlled by the AFP, were enhanced by experience, as they were strongest for mated females responding to their mate's directed songs. We then measured neural activity using expression of the immediate early gene product ZENK, and found that social context and song familiarity differentially modulated the number of ZENK-expressing cells in telencephalic auditory areas. Specifically, the number of ZENK-expressing cells in the caudomedial mesopallium (CMM) was most affected by whether a song was directed or undirected, whereas the caudomedial nidopallium (NCM) was most affected by whether a song was familiar or unfamiliar. Together these data demonstrate that females detect and prefer the features of directed song and suggest that high-level auditory areas including the CMM are involved in this social perception.
Author Summary
Vocal communication in many species, including humans, is affected by social cues. In the zebra finch, for example, males make subtle changes to the length, tempo, and variability of their courtship songs (directed songs) relative to songs performed in isolation (undirected songs). Using a behavioral approach assay, we found that female zebra finches strongly prefer the sound of directed over undirected song. Interestingly, female preferences were influenced by the variability of note pitch, showing stronger preferences for directed songs when they were less variable in pitch than the undirected songs. Pitch variability is controlled by a forebrain–basal ganglia circuit, which may represent a neural substrate on which selection acts to shape behavior. Preference for directed song was also increased when the singer was familiar to the listener, suggesting that song preferences are enhanced by experience. Based on the expression of an immediate early gene associated with memory formation and plasticity, we found that two high-level auditory areas were differentially responsive to the category of song females heard, with one area responding to whether songs were directed or undirected, and a second area to whether songs were familiar or unfamiliar. Together, these data demonstrate that females detect and prefer the male's changed performance during courtship singing and suggest that neurons in high-level auditory areas are involved in this social perception.
Female songbirds are attentive to subtle changes in male song, particularly to the variability of pitch. High-level auditory areas may generate females' behavioral preferences.
PMCID: PMC2267820  PMID: 18351801
3.  Song in the cold is ‘hot’: memory of and preference for sexual signals perceived under thermal challenge 
Biology Letters  2012;8(5):751-753.
The environmental conditions under which signals are perceived can affect receiver responses. Many songbird populations produce a song chorus at dawn, when, in cold habitats, they would experience thermal challenge. We recorded temperature and the song activity of Lincoln's sparrows (Melospiza lincolnii) on a high-elevation meadow, and determined that song behaviour is concentrated around the coldest time of the day, at dawn. We hypothesized that this is because male song in the cold is more attractive to females than song in the warm. To test this, we exposed laboratory-housed Lincoln's sparrow females to songs at 1°C and 16°C, which they naturally experience in the wild. Females spent 40 per cent more time close to the speaker during playback at 1°C than at 16°C. When tested at 16°C 1–2 days later, females biased their movement towards the speaker playing songs previously heard at 1°C over 16°C. Thus, female Lincoln's sparrows remembered and affiliated with songs they heard under thermal challenge, indicating that the thermal environment can affect the attractiveness of a sexual signal.
PMCID: PMC3441005  PMID: 22809726
dawn chorus; temperature; Melospiza lincolnii; mate choice; songbird
4.  Song Competition Affects Monoamine Levels in Sensory and Motor Forebrain Regions of Male Lincoln's Sparrows (Melospiza lincolnii) 
PLoS ONE  2013;8(3):e59857.
Male animals often change their behavior in response to the level of competition for mates. Male Lincoln's sparrows (Melospiza lincolnii) modulate their competitive singing over the period of a week as a function of the level of challenge associated with competitors' songs. Differences in song challenge and associated shifts in competitive state should be accompanied by neural changes, potentially in regions that regulate perception and song production. The monoamines mediate neural plasticity in response to environmental cues to achieve shifts in behavioral state. Therefore, using high pressure liquid chromatography with electrochemical detection, we compared levels of monoamines and their metabolites from male Lincoln's sparrows exposed to songs categorized as more or less challenging. We compared levels of norepinephrine and its principal metabolite in two perceptual regions of the auditory telencephalon, the caudomedial nidopallium and the caudomedial mesopallium (CMM), because this chemical is implicated in modulating auditory sensitivity to song. We also measured the levels of dopamine and its principal metabolite in two song control nuclei, area X and the robust nucleus of the arcopallium (RA), because dopamine is implicated in regulating song output. We measured the levels of serotonin and its principal metabolite in all four brain regions because this monoamine is implicated in perception and behavioral output and is found throughout the avian forebrain. After controlling for recent singing, we found that males exposed to more challenging song had higher levels of norepinephrine metabolite in the CMM and lower levels of serotonin in the RA. Collectively, these findings are consistent with norepinephrine in perceptual brain regions and serotonin in song control regions contributing to neuroplasticity that underlies socially-induced changes in behavioral state.
PMCID: PMC3608548  PMID: 23555809
5.  The Effect of Polychlorinated Biphenyls on the Song of Two Passerine Species 
PLoS ONE  2013;8(9):e73471.
Polychlorinated biphenyls (PCBs) are synthetic chemical pollutants with demonstrated detrimental toxic and developmental effects on humans and wildlife. Laboratory studies suggest that PCBs influence behavior due to their effects on endocrine and neurological systems, yet little is known about the behavioral consequences of sublethal PCB exposure in the field. Additionally, specific PCB congener data (in contrast to total PCB load) is necessary to understand the possible effects of PCBs in living organisms since number and position of chlorine substitution in a PCB molecule dictates the toxicity and chemical fate of individual PCB congeners. We non-lethally investigated total PCB loads, congener specific PCB profiles, and songs of black-capped chickadees (Poecile atricapillus) and song sparrows (Melospiza melodia) along a historical PCB gradient at the Hudson River in New York State. Our results indicate that black-capped chickadees and song sparrows have higher total blood PCBs in regions with higher historic PCB contamination. The two bird species varied substantially in their congener-specific PCB profiles; within sites, song sparrows showed a significantly higher proportion of lower chlorinated PCBs, while black-capped chickadees had higher proportions of highly chlorinated PCBs. In areas of PCB pollution, the species-specific identity signal in black-capped chickadee song varied significantly, while variation in song sparrow trill performance was best predicted by the mono-ortho PCB load. Thus, PCBs may affect song production, an important component of communication in birds. In conclusion, we suggest that the ramifications of changes in song quality for bird populations may extend the toxic effects of environmental PCB pollution.
PMCID: PMC3776824  PMID: 24058475
6.  Physically Challenging Song Traits, Male Quality, and Reproductive Success in House Wrens 
PLoS ONE  2013;8(3):e59208.
Physically challenging signals are likely to honestly indicate signaler quality. In trilled bird song two physically challenging parameters are vocal deviation (the speed of sound frequency modulation) and trill consistency (how precisely syllables are repeated). As predicted, in several species, they correlate with male quality, are preferred by females, and/or function in male-male signaling. Species may experience different selective pressures on their songs, however; for instance, there may be opposing selection between song complexity and song performance difficulty, such that in species where song complexity is strongly selected, there may not be strong selection on performance-based traits. I tested whether vocal deviation and trill consistency are signals of male quality in house wrens (Troglodytes aedon), a species with complex song structure. Males’ singing ability did not correlate with male quality, except that older males sang with higher trill consistency, and males with more consistent trills responded more aggressively to playback (although a previous study found no effect of stimulus trill consistency on males’ responses to playback). Males singing more challenging songs did not gain in polygyny, extra-pair paternity, or annual reproductive success. Moreover, none of the standard male quality measures I investigated correlated with mating or reproductive success. I conclude that vocal deviation and trill consistency do not signal male quality in this species.
PMCID: PMC3602011  PMID: 23527137
7.  Quality of song learning affects female response to male bird song. 
Bird song is unusual as a sexually selected trait because its expression depends on learning as well as genetic and other environmental factors. Prior work has demonstrated that males who are deprived of the opportunity to learn produce songs that function little if at all in male-female interactions. We asked whether more subtle variation in male song-learning abilities influences female response to song. Using a copulation solicitation assay, we measured the response of female song sparrows (Melospiza melodia) to songs of laboratory-reared males that differed in the amount of learned versus invented material that they included and in the degree to which learned material accurately matched the model from which it was copied. Females responded significantly more to songs that had been learned better, by either measure. Females did not discriminate between the best-learned songs of laboratory-reared males and songs of wild males used as models during learning. These results provide, to our knowledge, a first experimental demonstration that variation in learning abilities among males plays a functionally important part in the expression of a sexually selected trait, and further provide support for the hypothesis that song functions as an indicator of male quality because it reflects variation in response to early developmental stress.
PMCID: PMC1691117  PMID: 12350258
8.  Responses to song playback vary with the vocal performance of both signal senders and receivers 
Mating signals of many animal species are difficult to produce and thus should indicate signaler quality. Growing evidence suggests that receivers modulate their behaviour in response to signals with varying performance levels, although little is known about if and how responses are affected by receiver attributes. To explore this topic we conducted two experiments with swamp sparrows, Melospiza georgiana, in which we challenged territorial males with playback of songs with trill rates that were natural, digitally reduced, or digitally elevated (control-, low- and high-performance stimuli, respectively). In our first experiment, we found that males responded more aggressively to control songs than to low-performance stimuli, that low-performance stimuli with the most severe trill-rate reductions elicited the weakest aggressive responses, and that the subjects' own trill rates predicted aggressive responses. In our second experiment, we found that male responses to high-performance stimuli varied significantly, in ways predicted by two factors: the degree to which we had elevated stimulus performance levels of high-performance stimuli, and subjects' own vocal performance levels. Specifically, males were less aggressive towards stimuli for which we had elevated performance levels to higher degrees, and subject males with higher vocal performances themselves responded more aggressively. These findings together offer a novel illustration of how responses to aggressive signals may rely not just on signal attributes, but also on attributes of responding animals themselves.
PMCID: PMC3757973  PMID: 23945685
sexual selection; competition; performance; soft song; individual differences
9.  Independent effects of song quality and experience with photostimulation on expression of the immediate, early gene ZENK (EGR-1) in the auditory telencephalon of female European starlings 
Developmental neurobiology  2009;69(6):339-349.
Age influences behavioral decisions such as reproductive timing and effort. In photoperiodic species, such age effects may be mediated, in part, by the individual's age-accrued experience with photostimulation. In female European starlings (Sturnus vulgaris) that do not differ in age, experimental manipulation of photostimulation experience (photoexperience) affects hypothalamic, pituitary, and gonadal activity associated with reproductive development. Does photoexperience also affect activity in forebrain regions involved in processing a social cue, the song of males, which can influence mate choice and reproductive timing in females? Female starlings prefer long songs over short songs in a mate-choice context, and, like that in other songbird species, their auditory telencephalon plays a major role in processing these signals. We manipulated the photoexperience of female starlings, photostimulated them, briefly exposed them to either long or short songs, and quantified the expression of the immediate-early gene ZENK (EGR-1) in the caudomedial nidopallium as a measure of activity in the auditory telencephalon. Using an information theoretic approach, we found higher ZENK immunoreactivity in females with prior photostimulation experience than in females experiencing photostimulation for the first time. We also found that long songs elicited greater ZENK immunoreactivity than short song did. We did not find an effect of the interaction between photoexperience and song length, suggesting that photoexperience does not affect forebrain ZENK-responsiveness to song quality. Thus, photoexperience affects activity in an area of the forebrain that processes social signals, an effect that we hypothesize mediates, in part, the effects of age on reproductive decisions in photoperiodic songbirds.
PMCID: PMC2872984  PMID: 19224564
age and aging; Akaike Information Criterion (AIC); birdsong; experience; photoperiodism
10.  Ecological Validity and the Study of Procedural and Episodic Memory Function in Songbirds 
Cognitive sciences  2010;5(1):1-20.
Memory in songbirds, from song learning, production, and recognition to that for locations in complex environments, has led to the attractiveness of these animals as model systems for the changes occurring within and between neurons that lead to relevant modifications in behavior. Zebra finches (Taeniopygia guttata) in particular are excellent models attributable to their ability to readily perform the above-mentioned, ecologically relevant memories in the laboratories, and the ease with which these stereotyped behaviors can be manipulated and measured. This review centers on the independent functioning of and possible interactions between two primary memory systems in songbirds: those important for song or “procedural” memories, as well as those for place, such as food location, a “spatial” or “episodic-like” memory. Work over several decades has formed a relatively comprehensive understanding of the behavioral changes, neural substrates, and plasticity central to procedural memory (song learning and production) function in birds. However, few studies have examined spatial memory ability in those that do not store and retrieve caches, orient some distance away from and back to a home loft, or are not brood parasites. Zebra finches offer a rather unique advantage in this study of memory function and the interaction of memory systems: they do not store food, and are closed-ended song learners, biparental, not territorial, and non-migratory. Thus, their memory for song is not necessarily intertwined with that for time (of year) or location, as in a bird that learns a new song each breeding season, migrates to a particular breeding ground, or forgoes song and reproductive behavior in periods of food scarcity. Episodic-like memory in zebra finches is controlled by the hippocampus, and damage to this region, as in rodents and humans, compromises the ability to learn and/or remember particular spatial locations. In male zebra finches, hippocampal damage causes no appreciable, concurrent deficit in song learning or recognition. Interestingly, in females, while lesions do not disrupt a normal preference for conspecific over heterospecific songs, they do seem to abolish the preference for tutor song versus other novel, conspecific songs. It is therefore exciting to hypothesize a potential overlap between these memory systems. Support for this is provided by data from several anatomical, functional, and behavioral studies, chief among these that cells within the hippocampus show selectivity to conspecific but not other song stimuli and mate versus non-mate calls, and that several afferent and efferent projections to/of the hippocampus suggest a modulatory role for hippocampal neurons in song behavior. Specifically, we suggest that the hippocampus in zebra finches plays a role in “episodic-like characteristics of song perception,” making these birds exceptional models for examining functional overlaps among memory systems central to discrete, ecologically relevant behaviors.
PMCID: PMC3730287  PMID: 23914282
zebra finch; song learning; song perception; caudomedial nidopallium; hippocampus
11.  Sexual equality in zebra finch song preference: evidence for a dissociation between song recognition and production learning. 
Song in oscine birds is a culturally inherited mating signal and sexually dimorphic. From differences in song production learning, sex differences in song recognition learning have been inferred but rarely put to a stringent test. In zebra finches, Taeniopygia guttata, females never sing and the species has one of the greatest neuroanatomical differences in song-related brain nuclei reported for songbirds. Preference tests with sibling groups for which exposure to song had been identical during the sensitive phase for song learning in males, revealed equally strong influence of the tutor's song (here the father) on males' and females' adult song preferences. Both sexes significantly preferred the father's over unfamiliar song when having free control over exposure to playbacks via an operant task. The sibling comparisons suggest that this preference developed independently of the song's absolute quality: variation between siblings was as great as between nests. The results show that early exposure has an equally strong influence on males' and females' song preferences despite the sexual asymmetry in song production learning. This suggests that the trajectory for song recognition learning is independent of the one for song production learning.
PMCID: PMC1690953  PMID: 11934365
12.  Non-Breeding Song Rate Reflects Nutritional Condition Rather than Body Condition 
PLoS ONE  2012;7(5):e36547.
Numerous studies have focused on song in songbirds as a signal involved in mate choice and intrasexual competition. It is expected that song traits such as song rate reflect individual quality by being dependent on energetic state or condition. While seasonal variation in bird song (i.e., breeding versus non-breeding song) and its neural substrate have received a fair amount of attention, the function and information content of song outside the breeding season is generally much less understood. Furthermore, typically only measures of condition involving body mass are examined with respect to song rate. Studies investigating a potential relationship between song rate and other indicators of condition, such as physiological measures of nutritional condition, are scant. In this study, we examined whether non-breeding song rate in male European starlings (Sturnus vulgaris) reflects plasma metabolite levels (high-density lipoproteins (HDL), albumin, triglycerides and cholesterol) and/or body mass. Song rate was significantly positively related to a principal component representing primarily HDL, albumin and cholesterol (and to a lesser degree plasma triglyceride levels). There was only a trend toward a significant positive correlation between song rate and body mass, and no significant correlation between body mass and the abovementioned principal component. Therefore, our results indicate that nutritional condition and body mass represent different aspects of condition, and that song rate reflects nutritional rather than body condition. Additionally, we also found that intra-individual song rate consistency (though not song rate itself) was significantly positively related to lutein levels, but not to body mass or nutritional condition. Together our results suggest that the relation between physiological measures of nutritional condition and song rate, as well as other signals, may present an interesting line of future research, both inside and outside the breeding season.
PMCID: PMC3348915  PMID: 22590563
13.  Mate Choice in Adult Female Bengalese Finches: Females Express Consistent Preferences for Individual Males and Prefer Female-Directed Song Performances 
PLoS ONE  2014;9(2):e89438.
In the process of mate selection by female songbirds, male suitors advertise their quality through reproductive displays in which song plays an important role. Females evaluate the quality of each signal and the associated male, and the results of that evaluation guide expression of selective courtship displays. Some studies reveal broad agreement among females in their preferences for specific signal characteristics, indicating that those features are especially salient in female mate choice. Other studies reveal that females differ in their preference for specific characteristics, indicating that in those cases female evaluation of signal quality is influenced by factors other than simply the physical properties of the signal. Thus, both the physical properties of male signals and specific traits of female signal evaluation can impact female mate choice. Here, we characterized the mate preferences of female Bengalese finches. We found that calls and copulation solicitation displays are equally reliable indicators of female preference. In response to songs from an array of males, each female expressed an individual-specific song preference, and those preferences were consistent across tests spanning many months. Across a population of females, songs of some males were more commonly preferred than others, and females preferred female-directed songs more than undirected songs, suggesting that some song features are broadly attractive. Preferences were indistinguishable for females that did or did not have social experience with the singers, indicating that female preference is strongly directed by song features rather than experiences associated with the singer. Analysis of song properties revealed several candidate parameters that may influence female evaluation. In an initial investigation of those parameters, females could be very selective for one song feature yet not selective for another. Therefore, multiple song parameters are evaluated independently. Together these findings reveal the nature of signal evaluation and mate choice in this species.
PMCID: PMC3928452  PMID: 24558501
14.  D1-like dopamine receptor density in nuclei involved in social behavior correlates with song in a context-dependent fashion in male European starlings 
Neuroscience  2009;159(3):962-973.
Research in songbirds shows that singing behavior is regulated by both brain areas involved in vocal behavior as well as those involved in social behavior. Interestingly, the precise role of these regions in song can vary as a function of the social, environmental and breeding context. To date, little is known about the neurotransmitters underlying such context-dependent regulation of song. Dopamine (DA) modulates highly motivated, goal-directed behaviors (including sexually motivated song) and emerging data implicate DA in the context-dependent regulation of singing behavior. This study was performed to begin to examine whether differences in DA receptors may underlie, in part, context-dependent differences in song production. We used autoradiographic procedures to label D1-like and D2-like DA receptors to examine the relationship between DA receptor density and singing behavior in multiple contexts in male European starlings (Sturnus vulgaris). Within a breeding context (when testosterone (T) was high), D1-like receptor density in the medial preoptic nucleus (POM) and midbrain central gray (GCt) negatively correlated with song used to attract a female. Additionally in this context, D1-like receptor density in POM, GCt, medial bed nucleus of the stria terminalis (BSTm), and lateral septum (LS) negatively correlated with song likely used to defend a nestbox. In contrast, in a non-breeding context (when T was low), D1-like receptor density in POM and LS positively correlated with song used to maintain social flocks. No relationships were identified between song in any context and D2-like receptor densities. Differences in the brain regions and directional relationships between D1-like receptor binding and song suggest that dopaminergic systems play a region and context-specific role in song. These data also suggest that individual variation in singing behavior may, in part, be explained by individual differences in D1-like receptor density in brain regions implicated in social behavior.
PMCID: PMC2667808  PMID: 19356680
medial preoptic nucleus; midbrain central gray; medial bed nucleus of the stria terminalis; lateral septum; songbird; birdsong
15.  Swamp sparrows modulate vocal performance in an aggressive context 
Biology Letters  2008;5(2):163-165.
Vocal performance refers to the proficiency with which a bird sings songs that are challenging to produce, and can be measured in simple trilled songs by their deviation from an upper bound regression of frequency bandwidth on trill rate. Here, we show that male swamp sparrows (Melospiza georgiana) increase the vocal performance of individual song types in aggressive contexts by increasing both the trill rate and frequency bandwidth. These results are the first to demonstrate flexible modulation by songbirds of this aspect of vocal performance and are consistent with this signal feature having a role in aggressive communication.
PMCID: PMC2665810  PMID: 19087921
bird song; aggressive signalling; vocal performance
16.  Song types and their structural features are associated with specific contexts in the banded wren 
Animal behaviour  2005;70(4):921-935.
We studied the use of song types and their acoustic features in different social contexts in the banded wren (Thryothorus pleurostictus), a resident tropical songbird in which males possess about 20 distinctive song types varying in duration, bandwidth, note composition, and trill structure. We recorded six focal males intensively for four days each while we observed context information such as during versus after dawn chorus, presence of the female, counter-versus solo-singing, location at the edge versus centre of the territory, and proximity to the nest. All males used at least some song types differentially during each of these pairs of alternative contexts. Males also preferentially used the song types they shared with a given neighbour when interacting with that bird. Songs delivered during dawn chorus were significantly longer, wider in bandwidth, often compound (double songs), and more likely to contain a rattle or buzz and an up-sweeping trill, compared to songs delivered after dawn chorus. Similar features were also more commonly observed when birds were engaged in intense male-male interactions and boundary disputes after dawn chorus, especially when countersinging at the edge of the territory. The presence of the female caused the male to deliver song types with narrower whole-song and trill bandwidth and fewer rattles and buzzes, and song-type diversity and fraction of compound songs were higher when the female was present. Thus, in addition to using type matching and variations in song-type switching and diversity to signal different levels of aggressive intention, male banded wrens also select song types based on their acoustic structure in different social contexts.
PMCID: PMC1702368  PMID: 17173097
17.  Song and the song control pathway in the brain can develop independently of exposure to song in the sedge warbler. 
Previous studies have shown that female sedge warblers choose to mate with males that have more complex songs, and sexual selection has driven the evolution of both song complexity and the size of the major song control area (HVc) in the brain. In songbirds, learning from conspecifics plays a major role in song development and this study investigates the effects of isolation and exposure to song on song structure and the underlying song control system. Sibling pairs of hand-reared nestling sedge warblers were reared to sexual maturity under two conditions. Siblings in one group were reared individually in acoustic isolation in separate soundproof chambers. In the other group, siblings were reared together in an aviary with playback of recorded songs. The following spring, analysis of songs revealed that siblings reared in acoustic isolation produced normal song structures, including larger syllable repertoires than those exposed to song. We found no significant differences in the volumes of HVc, nucleus robustus archistnatalis, the lateral portion of the magnocellular nucleus and the density of dendritic spines between the two groups. Males exceeded females in all these measures, and also had a larger telencephalon. Our experiments show that complex song, sexual dimorphism in brain structure, and the size of song nuclei can all develop independently of exposure to song. These findings have important implications for how sexual selection can operate upon a complex male trait such as song and how it may also shape the more general evolution of brain structure in songbirds.
PMCID: PMC1691192  PMID: 12573065
18.  Do females invest more into eggs when males sing more attractively? Postmating sexual selection strategies in a monogamous reed passerine 
Ecology and Evolution  2014;4(8):1328-1339.
Maternal investment can play an important role for offspring fitness, especially in birds, as females have to provide their eggs with all the necessary nutrients for the development of the embryo. It is known that this type of maternal investment can be influenced by the quality of the male partner. In this study, we first verify that male song is important in the mate choice of female Eurasian reed warblers, as males mate faster when their singing is more complex. Furthermore, female egg investment varies in relation to male song characteristics. Interestingly, clutch size, egg weight, or size, which can be considered as an high-cost investment, is not influenced by male song characteristics, whereas comparably low-cost investment types like investment into diverse egg components are adjusted to male song characteristics. In line with this, our results suggest that female allocation rules depend on investment type as well as song characteristics. For example, egg white lysozyme is positively correlated with male song complexity. In contrast, a negative correlation exists between-song speed and syllable repetitiveness and egg yolk weight as well as egg yolk testosterone concentration. Thus, our results suggest that female egg investment is related to male song performance in several aspects, but female investment patterns regarding various egg compounds are not simply correlated.
PMCID: PMC4020692  PMID: 24834329
Acrocephalus scirpaceus; egg yolk testosterone; lysozyme; male quality; song
19.  Vocal performance influences male receiver response in the banded wren 
In a variety of songbirds the production of trilled song elements is constrained by a performance tradeoff between how fast a bird can repeat trill units (trill rate) and the range of frequencies each unit can span (frequency bandwidth). High-performance trills serve as an assessment signal for females, but little is known about the signal value of vocal performance for male receivers. We investigated the relationship between trill rate and frequency bandwidth in banded wren (Thryothorus pleurostictus) songs. Trilled song elements showed the same performance tradeoff found in other passerines and individuals differed in performance of some trill types. We tested the hypothesis that males of this species assess each other based on trill performance with a two-speaker experiment, in which territory owners were presented with alternating renditions of the same song type manipulated to differ in trill rate. Subjects were significantly more likely to approach the faster trill stimulus first. However, subjects that received trill types closer to the performance limit spent less time close to the fast speaker. Our results show that male banded wrens discriminate and respond differently to songs based on their vocal performance. Thus, performance of physically challenging songs may thus be important in intra- as well as intersexual assessment.
PMCID: PMC1634778  PMID: 16822751
bird song; sexual selection; trill rate; Thryothorus pleurostictus; performance limit; acoustic playback
20.  Vocal performance influences male receiver response in the banded wren 
In a variety of songbirds the production of trilled song elements is constrained by a performance tradeoff between how fast a bird can repeat trill units (trill rate) and the range of frequencies each unit can span (frequency bandwidth). High-performance trills serve as an assessment signal for females, but little is known about the signal value of vocal performance for male receivers. We investigated the relationship between trill rate and frequency bandwidth in banded wren (Thryothorus pleurostictus) songs. Trilled song elements showed the same performance tradeoff found in other passerines and individuals differed in performance of some trill types. We tested the hypothesis that males of this species assess each other based on trill performance with a two-speaker experiment, in which territory owners were presented with alternating renditions of the same song type manipulated to differ in trill rate. Subjects were significantly more likely to approach the faster trill stimulus first. However, subjects that received trill types closer to the performance limit spent less time close to the fast speaker. Our results show that male banded wrens discriminate and respond differently to songs based on their vocal performance. Thus, performance of physically challenging songs may be important in intra- as well as inter-sexual assessment.
PMCID: PMC1634778  PMID: 16822751
bird song; sexual selection; trill rate; Thryothorus pleurostictus; performance limit; acoustic playback
21.  Time course of changes in Gambel’s white-crowned sparrow song behavior following transitions in breeding condition 
Hormones and behavior  2008;55(1):217-227.
Seasonal changes in behavior and its underlying neural substrate are common across animal taxa. These changes are often triggered by steroid sex hormones. Song in seasonally breeding songbirds provides an excellent example of this phenomenon. In these species, dramatic seasonal changes mediated by testosterone and its metabolites occur in adult song behavior and in the neural circuitry controlling song. While song rate can quickly change in response to seasonal breeding cues, it is unknown how quickly other aspects of song change, particularly the stereotypy of song phonology and syntax. In this study we determined whether and how quickly song rate, phonology, and syntax change in response to breeding and non-breeding physiological cues. We asked these questions using Gambel’s white-crowned sparrows (Zonotrichia leucophrys gambelii), a closed-ended learner with well-characterized changes in the neural circuitry controlling song behavior. We exposed ten photosensitive sparrows to long-day photoperiod and implanted them with subcutaneous testosterone pellets (day 0) to simulate breeding conditions. We continuously recorded song and found that song rate increased quickly, reaching maximum around day 6. The stereotypy of song phonology changed more slowly, reaching maximum by day 10 or later. Song syntax changed minimally after day 6, the earliest time point examined. After 21 days, we transitioned five birds from breeding to non-breeding condition. Song rate declined precipitously. These results suggest that while song rate changes quickly, song phonology changes more slowly, generally following or in parallel with previously investigated changes in the neural substrate.
PMCID: PMC2648829  PMID: 19013173
Gambel’s white-crowned sparrow; Zonotrichia leucophrys gambelii; testosterone; birdsong; songbird; seasonal plasticity; photoperiod
22.  Birdsong: Is It Music to Their Ears? 
Since the time of Darwin, biologists have wondered whether birdsong and music may serve similar purposes or have the same evolutionary precursors. Most attempts to compare song with music have focused on the qualities of the sounds themselves, such as melody and rhythm. Song is a signal, however, and as such its meaning is tied inextricably to the response of the receiver. Imaging studies in humans have revealed that hearing music induces neural responses in the mesolimbic reward pathway. In this study, we tested whether the homologous pathway responds in songbirds exposed to conspecific song. We played male song to laboratory-housed white-throated sparrows, and immunolabeled the immediate early gene product Egr-1 in each region of the reward pathway that has a clear or putative homologue in humans. We found that the responses, and how well they mirrored those of humans listening to music, depended on sex and endocrine state. In females with breeding-typical plasma levels of estradiol, all of the regions of the mesolimbic reward pathway that respond to music in humans responded to song. In males, we saw responses in the amygdala but not the nucleus accumbens – similar to the pattern reported in humans listening to unpleasant music. The shared responses in the evolutionarily ancient mesolimbic reward system suggest that birdsong and music engage the same neuroaffective mechanisms in the intended listeners.
PMCID: PMC3508516  PMID: 23226128
Egr-1; mesolimbic reward system; reward; music; song; songbird
23.  Reward and Vocal Production: Song-associated Place Preference in Songbirds 
Physiology & behavior  2012;106(2):87-94.
Vocal production is crucial for successful social interactions in multiple species. Reward can strongly influence behavior; however, the extent to which reward systems influence vocal behavior is unknown. In songbirds, singing occurs in different contexts. It can be spontaneous and undirected (e.g., song produced alone or as part of a large flock) or directed towards a conspecific (e.g., song used to attract a mate or influence a competitor). In this study, we developed a conditioned place preference paradigm to measure reward associated with different types of singing behavior in two songbird species. Both male zebra finches and European starlings developed a preference for a chamber associated with production of undirected song, suggesting that the production of undirected song is tightly coupled to intrinsic reward. In contrast, neither starlings nor zebra finches developed a place preference in association with directed song; however, male starlings singing directed song that failed to attract a female developed a place aversion. Unsuccessful contact calling behavior was also associated with a place aversion. These findings suggest that directed vocal behavior is not tightly linked to intrinsic reward but may be externally reinforced by social interactions. Data across two species thus support the hypothesis that the production of undirected but not directed song is tightly coupled to intrinsic reward. This study is the first to identify song-associated reward and suggests that reward associated with vocal production differs depending upon the context in which communication occurs. The findings have implications for understanding what motivates animals to engage in social behaviors and ways in which distinct reward mechanisms function to direct socially appropriate behaviors.
PMCID: PMC3314114  PMID: 22285212
reward; reinforcement; motivation; social context; communication; birdsong
24.  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.
PMCID: PMC3889651  PMID: 24436508
Contextual variation in birdsong; Continuous singing style; Vocal consistency; Sound density; Alauda arvensis
25.  Vocal Communications and the Maintenance of Population Specific Songs in a Contact Zone 
PLoS ONE  2012;7(5):e35257.
Bird song has been hypothesized to play a role in several important aspects of the biology of songbirds, including the generation of taxonomic diversity by speciation; however, the role that song plays in speciation within this group may be dependent upon the ability of populations to maintain population specific songs or calls in the face of gene flow and external cultural influences. Here, in an exploratory study, we construct a spatially explicit model of population movement to examine the consequences of secondary contact of populations singing distinct songs. We concentrate on two broad questions: 1) will population specific songs be maintained in a contact zone or will they be replaced by shared song, and 2) what spatial patterns in the distribution of songs may result from contact? We examine the effects of multiple factors including song-based mating preferences and movement probabilities, oblique versus paternal learning of song, and both cultural and genetic mutations. We find a variety of conditions under which population specific songs can be maintained, particularly when females have preferences for their population specific songs, and we document many distinct patterns of song distribution within the contact zone, including clines, banding, and mosaics.
PMCID: PMC3344826  PMID: 22574116

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