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Learned bird song is influenced by inherited predispositions. The canary is a model system for the interaction of genes and learning on behaviour, especially because some strains have undergone artificial selection for song. In this study, roller canaries (bred for low-pitched songs) and border canaries (whose song is higher pitched, similar to the wild-type) were interbred and backcrossed to produce 58 males that sorted into seven genetically distinct groups. All males were tutored with the same set of songs, which included both low- and high-pitched syllables. Individuals were consistent within genetic groups but differed between groups in the proportion of low- versus high-pitched syllables they learned and sang. Both sex-linked and autosomal factors affected song learning and song production, in an additive manner. Dominant Z-chromosome factors facilitated high-pitched syllable learning and production, whereas the sex-linked alleles associated with the switch to low-pitched syllables under artificial selection were largely recessive. With respect to autosomal effects, the most surprising result is that males in the same genetic group had almost identical repertoires. This result challenges two common preconceptions: that genetic changes at different loci lead to distinct phenotypic changes, and that genetic predispositions affect learning in simple and general ways. Rather, different combinations of genetic changes can be associated with the same phenotypic effect; and predispositions can be remarkably specific, such as a tendency to learn and sing one song element rather than another.

Frequency modulation is a salient acoustic feature of birdsong. Its control is usually attributed to the activity of syringeal muscles, which affect the tension of the labia responsible for sound production. We use experimental and theoretical tools to test the hypothesis that for birds producing tonal sounds such as domestic canaries (Serinus canaria), frequency modulation is determined by both the syringeal tension and the air sac pressure. For different models, we describe the structure of the isofrequency curves, which are sets of parameters leading to sounds presenting the same fundamental frequencies. We show how their shapes determine the relative roles of syringeal tension and air sac pressure in frequency modulation. Finally, we report experiments that allow us to unveil the features of the isofrequency curves.

In the last two decades, maternally derived yolk androgens have been shown to significantly alter offspring development, and a number of these effects persist into adulthood. However, little is known about their underlying mechanisms. Mechanisms that have been suggested are changes in the endogenous androgen production post-hatching or changes in the sensitivity towards circulating androgens. We tested the effects of yolk testosterone on the plasma testosterone levels and the sensitivity to testosterone in 5months old male canaries that hatched from eggs that were either injected with testosterone (yT-males) or with a control solution (yC-males). Changes in sensitivity were investigated via the behavioural response to an experimental elevation of the plasma testosterone levels. We performed the experiment in fall (low endogenous testosterone production), focusing on testosterone dependent response traits (aggression and song). Before implantation, there was a non-significant trend that the plasma testosterone levels were lower in yT-males than in yC-males. Elevating the plasma testosterone concentrations increased aggressiveness, song bout length and similarity of repeated song elements (=consistency), with the latter likely being a consequence of testosterone-driven song crystallization. However, these effects were not different among yT- or yC-males in any of the parameters. Thus, our findings render it unlikely that changes in the sensitivity to testosterone post-hatching would form the main underlying mechanism of hormone-mediated maternal effects in birds. Further experiments are urgently needed in order to understand the nature of the phenotypic effects resulting from embryonic exposure to maternal yolk testosterone., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Understanding the different factors that may influence parasite virulence is of fundamental interest to ecologists and evolutionary biologists. It has recently been demonstrated that parasite virulence may occur partly through manipulation of host competitive ability. Differences in competitive ability associated with the social status (dominant or subordinate) of a host may determine the extent of this competition-mediated parasite virulence. We proposed that differences between subordinate and dominant birds in the physiological costs of infection may change depending on the level of competition in social groups. We observed flocks of domestic canaries to determine dominant or subordinate birds, and modified competition by providing restricted (high competition) or ad libitum food (low competition). Entire flocks were then infected with either the avian malaria parasite, Plasmodium relictum or a control. Contrary to our predictions we found that the level of competition had no effect on the outcome of infection for dominant or subordinate birds. We found that dominant birds appeared to suffer greater infection mediated morbidity in both dietary treatments, with a higher and more sustained reduction in haematocrit, and higher parasitaemia, than subordinates. Our results show that dominance status in birds can certainly alter parasite virulence, though the links between food availability, competition, nutrition and virulence are likely to be complex and multifaceted., (Copyright © 2013. Published by Elsevier Inc.)

Steroid hormones regulate multiple but distinct aspects of social behaviors. Testosterone (T) has multiple effects on learned courtship song in that it regulates both the motivation to sing in a particular social context as well as the quality of song produced. The neural substrate(s) where T acts to regulate the motivation to sing as opposed to other aspects of song has not been definitively characterized. We show here that T implants in the medial preoptic nucleus (POM) of castrated male canaries (Serinus canaria) increase song rate but do not enhance acoustic features such as song stereotypy compared with birds receiving peripheral T that can act globally throughout the brain. Strikingly, T action in the POM increased song control nuclei volume, consistent with the hypothesis that singing activity induces neuroplasticity in the song control system independent of T acting in these nuclei. When presented with a female canary, POM-T birds copulated at a rate comparable to birds receiving systemic T but produced fewer calls and songs in her presence. Thus, POM is a key site where T acts to activate copulation and increase song rate, an appetitive sexual behavior in songbirds, but T action in other areas of the brain or periphery (e.g., HVC, dopaminergic cell groups, or the syrinx) is required to enhance the quality of song (i.e., stereotypy) as well as regulate context-specific vocalizations. These results have broad implications for research concerning how steroids act at multiple brain loci to regulate distinct sociosexual behaviors and the associated neuroplasticity.

There is an increasing understanding of the context-dependent nature of parasite virulence. Variation in parasite virulence can occur when infected individuals compete with conspecifics that vary in infection status; virulence may be higher when competing with uninfected competitors. In vertebrates with social hierarchies, we propose that these competition-mediated costs of infection may also vary with social status. Dominant individuals have greater competitive ability than competing subordinates, and consequently may pay a lower prevalence-mediated cost of infection. In this study we investigated whether costs of malarial infection were affected by the occurrence of the parasite in competitors and social status in domestic canaries (Serinus canaria). We predicted that infected subordinates competing with non-infected dominants would pay higher costs than infected subordinates competing with infected dominants. We also predicted that these occurrence-mediated costs of infection would be ameliorated in infected dominant birds. We found that social status and the occurrence of parasites in competitors significantly interacted to change haematocrit in infected birds. Namely, subordinate and dominant infected birds differed in haematocrit depending on the infection status of their competitors. However, in contrast to our prediction, dominants fared better with infected subordinates, whereas subordinates fared better with uninfected dominants. Moreover, we found additional effects of parasite occurrence on mortality in canaries. Ultimately, we provide evidence for costs of parasitism mediated by social rank and the occurrence of parasites in competitors in a vertebrate species. This has important implications for our understanding of the evolutionary processes that shape parasite virulence and group living., (Copyright © 2013 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

The nature of telencephalic control over premotor and motor circuits is debated. Hypotheses range from complete usurping of downstream circuitry to highly interactive mechanisms of control. We show theoretically and experimentally, that telencephalic song motor control in canaries is consistent with a highly interactive strategy. As predicted from a theoretical model of respiratory control, mild cooling of a forebrain nucleus (HVC) led to song stretching, but further cooling caused progressive restructuring of song, consistent with the hypothesis that respiratory gestures are subharmonic responses to a timescale present in the output of HVC. This interaction between a life-sustaining motor function (respiration) and telencephalic song motor control suggests a more general mechanism of how nonlinear integration of evolutionarily new brain structures into existing circuitry gives rise to diverse, new behavior.

It is well established that auditory forebrain regions of oscine birds are essential for the encoding of species-typical songs and are, therefore, vital for recognition of song during sociosexual interactions. Regions such as the caudal medial nidopallium (NCM) and the caudal medial mesopallium (CMM) are involved in perceptual processing of song and the formation of auditory memories. There is an additional telencephalic nucleus, however, that has also been implicated in species recognition. This nucleus is HVC, a prominent nucleus that sits at the apex of the song system, and is well known for its critical role in song learning and song production in male songbirds. Here, we explore the functional relationship between auditory forebrain regions (i.e., NCM and CMM) and HVC in female canaries (Serinus canaria). We lesion HVC and examine immediate early gene responses to conspecific song presentation within CMM and NCM to explore whether HVC can modulate auditory responses within these forebrain regions. Our results reveal robust deficits in ZENK-ir in CMM and NCM of HVC-lesioned females when compared with control- and sham-lesioned females, indicating that functional connections exists between HVC and NCM/CMM. Although these connected regions have been implicated in song learning and production in males, they likely serve distinct functions in female songbirds that face the task of song recognition rather than song production. Identifying functional connections between HVC and auditory regions involved in song perception is an essential step toward developing a comprehensive understanding of the neural basis of song recognition., (Copyright © 2012 Wiley Periodicals, Inc.)

Bird songs range in form from the simple notes of a Chipping Sparrow to the rich performance of the nightingale. Non-adjacent correlations can be found in the syntax of some birdsongs, indicating that the choice of what to sing next is determined not only by the current syllable, but also by previous syllables sung. Here we examine the song of the domesticated canary, a complex singer whose song consists of syllables, grouped into phrases that are arranged in flexible sequences. Phrases are defined by a fundamental time-scale that is independent of the underlying syllable duration. We show that the ordering of phrases is governed by long-range rules: the choice of what phrase to sing next in a given context depends on the history of the song, and for some syllables, highly specific rules produce correlations in song over timescales of up to ten seconds. The neural basis of these long-range correlations may provide insight into how complex behaviors are assembled from more elementary, stereotyped modules.

The preference of female songbirds for particular traits in the songs of courting males has received considerable attention, but the relationship of preferred traits to male quality is poorly understood. Female domestic canaries (Serinus canaria, Linnaeus) preferentially solicit copulation with males that sing special high repetition rate, wide-band, multi-note syllables, called 'sexy' or A-syllables. Syllables are separated by minibreaths but each note is produced by pulsatile expiration, allowing high repetition rates and long duration phrases. The wide bandwidth is achieved by including two notes produced sequentially on opposite sides of the syrinx, in which the left and right sides are specialized for low or high frequencies, respectively. The emphasis of low frequencies is facilitated by a positive relationship between syllable repetition rate and the bandwidth of the fundamental frequency of notes sung by the left syrinx, such that bandwidth increases with increasing syllable repetition rate. The temporal offset between notes prevents cheating by unilaterally singing a note on the left side with a low fundamental frequency and prominent higher harmonics. The syringeal and respiratory motor patterns by which sexy syllables are produced support the hypothesis that these syllables provide a sensitive vocal-auditory indicator of a male's performance limit for the rapid, precisely coordinated interhemispheric switching, which is essential for many sensory and motor processes involving specialized contributions from each cerebral hemisphere.

Song discrimination and recognition in songbird species have usually been studied by measuring responses to song playbacks. In female canaries, Serinus canaria, copulation solicitation displays (CSDs) are used as an index of female preferences, which are related to song recognition. Despite the fact that many studies underline the role of song syntax in this species, we observed that short segments of songs (a few seconds long) are enough for females to discriminate between conspecific and heterospecific songs, whereas such a short duration is not sufficient to identify the syntax rules. This suggests that other cues are salient for song recognition. In this experiment, we investigated the influence of amplitude modulation (AM) on the responses (CSDs) of female canaries to song playbacks. We used two groups of females: (1) raised in acoustic isolation and (2) raised in normal conditions. When adult, we tested their preferences for sexy phrases with different AMs. We broadcast three types of stimuli: (1) songs with natural canary AM, (2) songs with AM removed, or (3) song with wren Troglodytes troglodytes AM. Results indicate that female canaries prefer and have predispositions for a song type with the natural canary AM. Thus, this acoustic parameter is a salient cue for song attractiveness.

In many bird species with hatching asynchrony, yolk androgens increase across the laying sequence. This has been hypothesized to represent a compensatory mechanism for disadvantages of later-hatching chicks - via positive effects of yolk androgens on early competitiveness and growth. However, the costs and benefits of this compensatory strategy probably depend on environmental factors determining the survival chances of the chicks such as the food conditions, which should, therefore, influence maternal yolk androgen deposition. We studied the consequences of manipulated food conditions on the expected level of hatching asynchrony in canaries (Serinus canaria) assigning females to either a low (=LQ) or high quality (=HQ) diet. We measured the incubation behaviour (as incubation attendance) and the yolk androgen deposition in order to investigate whether and how females modulate hatching asynchrony in relation to the food conditions. Females on a HQ diet laid larger and heavier clutches, showed a stronger increase in yolk testosterone content towards the last-laid eggs, but did not alter their incubation attendance. Thus, females on a HQ diet seem to favour the survival of later hatching chicks, as indicated by their yolk testosterone deposition pattern. However, females on a HQ diet laid larger clutches and might need to compensate more in order to achieve a similar degree of hatching asynchrony than females on a LQ diet, given the lack of plasticity in incubation attendance. This suggests that canary females respond to food manipulations mainly via changes in clutch size rather than by altering the degree of hatching asynchrony., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Maternal effects occur when the phenotype of the mother affects the phenotype of their offspring. They are thought to have evolved to translate the environmental conditions experienced by the mother into adaptive phenotypic variation of the offspring. However, the integration of environmental cues allowing adaptive responses requires some form of plasticity that depends on the interaction of the maternal phenotype and her environment. In birds, maternal yolk hormones represent such a pathway for maternal effects, and their adaptive significance depends thus on the plasticity in maternal yolk hormone deposition. We studied sources of variation in yolk testosterone deposition, focusing on the often neglected contribution of the (partly heritable) maternal phenotype. We investigated consistency and heritability of yolk testosterone deposition in captive canaries of which the F(1) generation was raised in foster nests and analyzed the potential effects of the early developmental conditions. We found significant female consistency across years in egg mass, yolk mass and total amount of yolk testosterone but not in yolk testosterone concentrations. Females varied the yolk testosterone concentrations of their eggs across years mainly via changes in yolk mass. The heritable variation in egg mass, yolk mass and amount of yolk testosterone but not yolk testosterone concentrations was within the range of previous studies, but not significantly different from zero. Finally, the growth of the daughters as nestling had a significant effect on their yolk testosterone deposition at adulthood indicating the transgenerational potential for environmental effects - via the effects of yolk hormones on offspring development., (Copyright © 2011 Elsevier Inc. All rights reserved.)

In most temperate zone vertebrates, photoperiodic change plays the major role in the timing of seasonal breeding. However, direct environmental stimuli such as temperature, rainfall, or availability of food are thought to be important for fine-tuning breeding activities. Building on evidence from wild Island canaries (Serinus canaria), the authors had shown advancing effects of green vegetation on breeding under captive, short-day conditions. So far, the precise, sensory modalities of this stimulatory cue are unknown. Here the authors present new data that confirm advanced breeding activities in the presence of green vegetation and narrow its stimulatory components. They found that direct exposure of the birds to fresh green vegetation represented the strongest stimulus and advanced breeding by up to 2 months compared to controls. In contrast, access to artificial green vegetation, extracts from green vegetation, or olfactory components alone had no such effects. This is, to the best of the authors' knowledge, the first experiment that examines sensory components of an effective, supplementary, non-photoperiodic cue in a temperate zone species. The data suggest that in order to use non-photoperiodic information to time breeding, birds must be able to integrate and process multisensory stimuli. Single non-photoperiodic sensory cues are insufficient to affect the timing of seasonal breeding., (© 2011 The Author(s))

Auditory brainstem responses (ABRs) were recorded in adult budgerigars, canaries, and zebra finches in quiet and in three levels of white noise for tone stimuli between 1 and 4 kHz. Similar to behavioral results, masked ABR thresholds increased linearly with increasing noise levels. When the three species are considered together, ABR-derived CRs were higher than behavioral CRs by 18-23 dB between 2 and 4 kHz and by about 30 dB at 1 kHz. This study clarifies the utility of using ABRs for estimating masked auditory thresholds in natural environmental noises in species that cannot be tested behaviorally., (© 2011 Acoustical Society of America)

In songbirds, neurons in the song nucleus HVC exhibit a striking example of selective auditory response, firing more to playback of the bird's own song (BOS) than to conspecific songs. This song selectivity has been found in various songbird species, both those that sing a single individual-specific song as well as those, such as the canary, in which both song structure and individual-identity encoding in song is more complex. In the present study, we investigated how the BOS is represented in the HVC of anesthetized long-day canaries by using temporal and spectral variants of the BOS stimulus. We addressed the question of how selective HVC neurons were by quantifying the number of song elements, called phrases, that evoked auditory responses. Phrases that were individual-specific or that were frequently delivered in an individual's songs did not drive HVC neurons to a greater degree than others. Reordering phrases or altering their acoustic structure caused a decrease in the auditory responsiveness of HVC neurons. This sensitivity to the spectral and temporal features of the BOS involved neurons that failed to respond to BOS variants or were driven by a reduced number of phrases, as well as neurons whose auditory responsiveness extended beyond the features of the individual's song, responding to phrases that were not sung by the bird itself. Therefore, the neural strategy by which BOS structure is represented in the canary HVC may require something other than a strict representation of the repertoire of song components. We suggest that the individual's song could be coded, at least in part, by an ensemble of broadly tuned neurons., (© 2011 IBRO. Published by Elsevier Ltd. All rights reserved.)

Many animals live in a communication network, an environment where individuals can obtain information about competitors or potential mates by observing interactions between conspecifics. In such an environment, interactants might benefit by changing their signalling behaviour in the presence of an audience. This audience effect seems widespread among species, has been observed during various types of interaction (e.g. intra-sexual vs. inter-sexual interaction) and varies according to the social context (e.g. gender, hierarchical or mating status of the audience). However, the way individuals might adapt their signalling behaviour to a combination of these factors remains poorly understood. To address this question, we studied how the presence of an audience affects the behaviour of male domestic canaries Serinus canaria during two types of interactions: (i) an extra-pair interaction and (ii) a male-male competition for food. Males were observed under three conditions: (a) in the absence of audience, (b) in the presence of their mate or (c) of a familiar female. Our results show that male domestic canaries minutely adapt their courting and agonistic behaviours to a combination of: (i) the type of interaction (extra-pair interaction/male-male competition), (ii) the social context (mate, familiar female or nobody in audience) and (iii) the behaviours of both the audience and the interactant. These results highlight the ability of animals to subtly adapt their behaviour to the social environment. This also raises questions about the cognitive foundations and evolution of these processes especially considering that canaries are known neither for having high cognitive abilities nor for being a typical example for the social intelligence hypothesis.

The brain of adult homeothermic vertebrates exhibits a higher degree of morphological neuroplasticity than previously thought, and this plasticity is especially prominent in birds. In particular, incorporation of new neurons is widespread throughout the adult avian forebrain, and the volumes of specific nuclei vary seasonally in a prominent manner. We review here work on steroid-dependent plasticity in birds, based on two cases: the medial preoptic nucleus (POM) of Japanese quail in relation to male sexual behavior, and nucleus HVC in canaries, which regulates song behavior. In male quail, POM volume changes seasonally, and in castrated subjects testosterone almost doubles POM volume within 2 weeks. Significant volume increases are, however, already observable after 1 day. Steroid receptor coactivator-1 is part of the mechanism mediating these effects. Increases in POM volume reflect changes in cell size or spacing and dendritic branching, but are not associated with an increase in neuron number. In contrast, seasonal changes in HVC volume reflect incorporation of newborn neurons in addition to changes in cell size and spacing. These are induced by treatments with exogenous testosterone or its metabolites. Expression of doublecortin, a microtubule-associated protein, is increased by testosterone in the HVC but not in the adjacent nidopallium, suggesting that neuron production in the subventricular zone, the birthplace of newborn neurons, is not affected. Together, these data illustrate the high degree of plasticity that extends into adulthood and is characteristic of avian brain structures. Many questions still remain concerning the regulation and specific function of this plasticity., (© 2010 The Authors. European Journal of Neuroscience © 2010 Federation of European Neuroscience Societies and Blackwell Publishing Ltd.)

In songbirds, the ontogeny of singing behavior shows strong parallels with human speech learning. As in humans, development of learned vocal behavior requires exposure to an acoustic model of species-typical vocalizations, and, subsequently, a sensorimotor practice period after which the vocalization is produced in a stereotyped manner. This requires mastering motor instructions driving the vocal organ and the respiratory system. Recently, it was shown that, in the case of canaries (Serinus canaria), the diverse syllables, constituting the song, are generated with air sac pressure patterns with characteristic shapes, remarkably, those belonging to a very specific mathematical family. Here, we treated juvenile canaries with testosterone at the onset of the sensorimotor practice period. This hormone exposure accelerated the development of song into stereotyped adultlike song. After 20 days of testosterone treatment, subsyringeal air sac pressure patterns of song resembled those produced by adults, while those of untreated control birds of the same age did not. Detailed temporal structure and modulation patterns emerged rapidly with testosterone treatment, and all previously identified categories of adult song were observed. This research shows that the known effect of testosterone on the neural circuits gives rise to the stereotyped categories of respiratory motor gestures. Extensive practice of these motor patterns during the sensorimotor phase is not required for their expression., (Copyright © 2010 Wiley Periodicals, Inc.)

Belgian Waterslager (BW) canaries have an inherited hearing loss due to missing and abnormal hair cells, but it is unclear whether the loss is congenital or developmental. We used auditory brainstem responses and scanning electron microscopy to describe the development of auditory sensitivity and hair cell abnormalities in BW and non-BW canaries. In both strains, adult ABR thresholds were higher than behavioral thresholds, but BW canaries exhibited higher thresholds than non-BW canaries across all frequencies. Immediately post-hatch, ABR thresholds and hair cell numbers were similar in both strains. Two weeks later, thresholds were significantly higher in BW canaries, and hair cell number progressively decreased as the birds aged. These data show that in BW canaries: the peripheral auditory system is functionally similar to non-BW canary from hatch to 2 weeks, ABR thresholds improve during this developmental period, actually becoming better than those of adults, but then worsen as the bird continues to age. Hair cell number and appearance is similar to non-BW canaries at hatch but progressively declines after 30 days of age. These data show that the hearing loss characteristic of BW canaries is, at least in part, developmental and is established by the time song learning begins.

Neuron recruitment has been implicated in morphological and functional plasticity in the adult brain. Whereas mammals restrict neuron recruitment specifically to two regions of known plasticity, the hippocampus and olfactory bulb, newborn neurons are found throughout the forebrain of adult songbirds. In order to study the area-specificity of the widespread proliferation and recruitment in the songbird brain, six adult male canaries received repetitive intraperitoneal injections of the mitotic marker BrdU (5-bromo-2-deoxyuridine) and were sacrificed after 24 hours to study proliferation or after 38 days to study recruitment. Migration and incorporation of new neurons was apparent throughout many but not all parts of the canary forebrain and was quantitatively related to mitotic levels in the most closely associated proliferative zones. Surprisingly, some areas of the vocal control system sensitive to plastic changes, such as nucleus higher vocal center (HVC) and area X, recruited similar numbers of new neurons as their surrounding brain tissues, employing no specific directional mechanisms. The distribution pattern in and around HVC could best be described by a random displacement model, where cells originating from the overlying lateral ventricle can move independently in any direction. Other plastic song control areas, such as the medial magnocellular nucleus of anterior nidopallium and the robust nucleus of arcopallium, were specifically avoided by migrating neurons, while migration toward the olfactory bulb showed high specificity, similar to the mammalian rostral migratory stream. Thus, different mechanisms appear to organize area-specific neuron recruitment in different recipients of the adult songbird brain, unrelated to global plasticity of brain regions., ((c) 2009 Wiley-Liss, Inc.)

In songbirds, experience of social and environmental cues during a discrete period after birth may dramatically influence song learning. In the canary, the ability to learn new songs is assumed to persist throughout life. The aim of the present study was to investigate whether social context could guide changes in adult song. Three groups of canaries were kept in different social and temporal conditions. Results showed that the multiple hierarchical levels of the canary song structure were affected by social environment: songs of males housed together for 2 years were more similar than those of males that spent the same time in individual cages in regard to acoustic parameters, syllable repertoire and repertoire of sequences of two-syllable types. However, social housing did not result in the emergence of a group-specific vocal signature within songs. In conclusion, these results suggested that under the influence of social factors, a copying process could allow adult canaries to adjust, at least in part, their songs to those of other individuals.

Many songbirds learn their songs early in life from a song model. In the absence of such a model, they develop an improvised song that often lacks the species-typical song structure. Open-ended learners, such as the domesticated canary, are able to modify their songs in adulthood, although the mechanisms that guide and time the song-learning process are still not fully understood. In a previous study, we showed that male domesticated canaries lacking an adult song model in their first year substantially change their song repertoire and composition when exposed to normally reared conspecifics in their second year. Here, we investigate song development in descendants of canaries that were raised and kept as a peer group without a song model. Such males represent tutors with abnormal song characteristics. Interestingly, the F(1) generation developed quite normal song structure, and when brought into an environment with normally raised canaries in their second year, they did not modify their songs substantially. These results suggest that contact with an adult song model early in life is crucial for song crystallization, but also that song development is at least partly guided by innate rules. They also question the existing classification of canaries as open-ended learners.

The solicitation behaviours performed by dependent young are under selection from the environment created by their parents, as well as wider ecological conditions. Here we show how mechanisms acting before hatching enable canary offspring to adapt their begging behaviour to a variable post-hatching world. Cross-fostering experiments revealed that canary nestling begging intensity is positively correlated with the provisioning level of their own parents (to foster chicks). When we experimentally increased food quality before and during egg laying, mothers showed higher faecal androgen levels and so did their nestlings, even when they were cross-fostered before hatching to be reared by foster mothers that had been exposed to a standard regime of food quality. Higher parental androgen levels were correlated with greater levels of post-hatching parental provisioning and (we have previously shown) increased faecal androgens in chicks were associated with greater begging intensity. We conclude that androgens mediate environmentally induced plasticity in the expression of both parental and offspring traits, which remain correlated as a result of prenatal effects, probably acting within the egg. Offspring can thus adapt their begging intensity to variable family and ecological environments.

During song production, oscine birds produce large air sac pressure pulses. During those pulses, energy is transferred to labia located at the juncture between the bronchii and the trachea, inducing the high frequency labial oscillations which are responsible for airflow modulations, i.e., the uttered sound. In order to generate diverse syllables, canaries (Serinus canaria) use a set of air sac pressure patterns with characteristic shapes. In this work we show that these different shapes can be approximated by the subharmonic solutions of a forced normal form. This simple model is built from identifying dynamical elements which allow to reproduce the shape of the pressure pattern corresponding to one syllable type. Remarkably, integrating that simple model for other parameters allows to recover the other pressure patterns used during song. Interpreting the diversity of these physiological gestures as subharmonic solutions of a simple nonlinear system allows us to account simultaneously for their morphological features as well as for the syllabic timing and suggests a strategy for the generation of complex motor patterns.

In songbirds, auditory neurons of the nucleus HVC respond selectively to a particular complex sound, the bird's own song (BOS). In the canary, this song selectivity did not exclude responses to conspecific songs. Here, we recorded single units in nucleus HVC of adult canaries to assess to what extent repertoire sharing among birds contributed to auditory responsiveness to birds' songs other than the BOS. Results indicated that song phrases driving auditory responses could differ from bird's own phrases suggesting that a subset of neurons were not strictly tuned to acoustic features of self-generated song components. In the canary, auditory representation of the BOSs might be more complex than that which has been described for birds with a small repertoire.

During acoustic communication, animals must attend to sounds from a particular source while simultaneously rejecting intrusion from other sources. One possible candidate mechanism for this process is the noradrenergic system. Noradrenaline is a neuromodulator that tunes sensory processing systems and regulates attention. We examined whether pharmacological degradation of the noradrenergic system using N-(2-chloroethyl)-N-2-bromobenzyl-amine hydrochloride (DSP-4) modifies processing of species-typical auditory signals in female canaries (Serinus canaria). We measured auditory responses to conspecific and heterospecific songs using ZENK protein expression within the caudomedial nidopallium (NCM) and the mesopallium caudomedial (CMM). Song-induced ZENK expression in these auditory forebrain areas is typically higher in birds exposed to conspecific songs as opposed to heterospecific songs. Our results reveal that this differential ZENK induction is abolished specifically within dNCM and CMM in female canaries treated with DSP-4. Furthermore, in DSP-4-treated birds, conspecific song-induced ZENK expression is significantly reduced when compared to saline-treated birds. This suggests that the noradrenergic system modifies auditory processing by enhancing neuronal responses to signals relevant to survival and reproduction rather than inhibiting neuronal responses to signals that are less relevant. Overall, our results reveal that noradrenaline plays a significant neuromodulatory role during the reception of species-typical communication signals., (Copyright 2008 S. Karger AG, Basel.)

The caudomedial nidopallium (NCM) is a telencephalic auditory area that is selectively activated by conspecific vocalizations in zebra finches and canaries. We recently demonstrated that temporal and spectral dynamics of auditory tuning in NCM differ between these species [1]. In order to determine whether these differences reflect recent experience, we exposed separate groups of each species and sex to different housing conditions. Adult birds were housed either in an aviary with conspecifics (NORM), with heterospecifics (canary subjects in a zebra finch aviary, and vice versa: (CROSS)), or in isolation (ISO) for 9 days prior to testing. We then recorded extracellular multi-unit electrophysiological responses to simple pure tone stimuli (250-5000 Hz) in awake birds from each group and analyzed auditory tuning width using methods from our earlier studies. Relative to NORM birds, tuning was narrower in CROSS birds, and wider in ISO birds. The trend was greater in canaries, especially females. The date of recording was also included as a covariate in ANCOVAs that analyzed a larger set of the canary data, including data from birds tested outside of the breeding season, and treated housing condition and sex as independent variables. These tests show that tuning width was narrower early in the year and broader later. This effect was most pronounced in CROSS males. The degree of the short-term neural plasticity described here differs across sexes and species, and may reflect differences in NCM's anatomical and functional organization related to species differences in song characteristics, adult plasticity and/or social factors. More generally, NCM tuning is labile and may be modulated by recent experience to reflect the auditory processing required for behavioral adaptation to the current acoustic, social or seasonal context.

The song of the domesticated canary (Serinus canaria) is one of the most widely used models to study the neural correlates of behavioural plasticity and the mechanisms of female mate choice. However, only few studies have described the song behaviour in detail and monitored their changes throughout the year, and these data are restricted to the "Waterslager" strain. Here, we studied the song characteristics of the male common domesticated canary at different times of the year, the spring breeding and autumnal non-breeding season, and monitored the birds' songs up to the following breeding season. During breeding, males have increased plasma levels of testosterone, and songs are on average longer and consist of fewer non-repeated syllable types compared to the non-breeding season. When subsequent seasons are compared, song duration and the proportion of non-repeated syllable types change seasonally but not across years. Repertoire size remains constant throughout seasons although syllable types are exchanged. Syllable carry-over is significantly higher from one breeding season to the next than between the breeding and non-breeding season. Further, the repertoire of the breeding season contains more potentially sexually attractive syllable types than that of the non-breeding season. These data show that overall song structure is retained throughout the year while seasonality occurs in the temporal pattern and in repertoire composition.

In temperate zone songbirds, such as canaries (Serinus canaria), seasonal variation in gonadal activity and behavior are associated with marked brain changes. These include gonadotropin-releasing hormone (GnRH) expression and the volume of brain areas controlling song production. Questions have been raised about the consistency of seasonal brain changes in canaries. Laboratory studies of the American singer strain raised doubts as to whether this strain exhibits a robust photoperiodic response along with changes in brain GnRH content, and studies of free-living canaries have failed to identify seasonal changes in volume of song control nuclei. We assessed differences in brain GnRH and the song control system associated with photoperiod-induced variation in reproductive state in Border canaries. We found that males and females maintained for 10 weeks on long days (14L:10D) regress their gonads, exhibit a decline in testosterone and initiate molt; a response consistent with the onset of absolute photorefractoriness (i.e., failed to respond to previously stimulating daylengths). All birds regained photosensitivity (i.e., exhibited gonadal response to stimulating daylengths) after experiencing short days (8L:16D) for 6 weeks. Furthermore, comparisons of birds in either a photosensitive, photostimulated, or photorefractory state revealed a marked increase in GnRH protein expression in the photosensitive and photostimulated birds over photorefractory birds. A similar variation was observed in the volume of key forebrain song nuclei. Thus, Border canaries demonstrate measurable neuroplasticity in response to photoperiodic manipulations. These data, along with previous work on other strains of canaries, indicate the presence of intra-specific variation in photoperiodically regulated neuroplasticity.

Nestling birds solicit food from their parents with vigorous begging displays, involving posturing, jostling and calling. In some species, such as canaries, begging is especially costly because it causes a trade off against nestling growth. Fitness costs of begging like this are predicted by evolutionary theory because they function to resolve conflicts of interest within the family over the provision of parental investment. However, the mechanism that links these costs with nestling behaviour remains unclear. In the present study, we determine if the relationships between nestling androgen levels, nestling begging intensities and nestling growth rates are consistent with the hypothesis that testosterone is responsible for the trade-off between begging and growth. We test this idea with a correlational study, using fecal androgens as a non-invasive method for assaying nestling androgen levels. Our results show that fecal androgen levels are positively correlated with nestling begging intensity, and reveal marked family differences in each trait. Furthermore, changes in fecal androgen levels between 5 and 8 days after hatching are positively associated with changes in nestling begging intensity, and negatively associated with nestling growth during this time. Although these correlational results support our predictions, we suggest that that experimental manipulations are now required to test the direct or indirect role of testosterone in mediating the trade-off between begging and growth.

The caudomedial nidopallium (NCM) of songbirds is a telencephalic area involved in the auditory processing and memorization of complex vocal communication signals. We used pure tone stimuli and multiunit electrophysiological recordings in awake birds to investigate whether the basic properties of song-responding circuits in NCM differ between canaries and zebra finches, two species whose songs are markedly different in their spectral and temporal organization. We found that the responses in zebra finch NCM are characterized by broad tuning and sustained responses that may facilitate the integration of zebra finch song syllables and call notes that are of long duration and have a broad harmonic structure. In contrast, we found that the responses in canary NCM show narrower tuning and less sustained responses over the time periods analyzed. These characteristics may contribute to enhanced processing of the narrow-band whistles, rapid trills, and steep frequency modulations that are prominent features of canary song. These species differences are much less pronounced in field L2, the direct thalamorecipient region that represents a preceding station in the central avian auditory pathway. NCM responses did not differ across sexes of either species, but field L2 did show wider tuning in zebra finch females relative to males. In sum, species differences in the response properties of NCM likely reflect selectivity for the acoustic elements of each species' vocal repertoire., ((c) 2007 Wiley Periodicals, Inc.)

The protein doublecortin (DCX) is expressed in post-mitotic migrating and differentiating neurons in the developing vertebrate brain and, as a part of the microtubule machinery, is required for neuronal migration. DCX expression is generally maximal during embryonic and early post-natal life but decreases markedly and almost disappears in older animals in parallel with the major decrease or cessation of neurogenesis. In several seasonally breeding songbird species such as canaries, the volume of several song control nuclei in the brain varies seasonally such that the largest nuclei are observed in the late spring and early summer. This variation is based on changes in cell size, dendritic branching, and, in nucleus HVC, on the incorporation of neurons newly born in adulthood. Because songbirds maintain an active neurogenesis and neuronal incorporation in their telencephalon throughout their lives, we investigated here the distribution of DCX-immunoreactive (ir) structures in the brain of adult male canaries. Densely stained DCX-ir cells were found exclusively in parts of the telencephalon that are known to incorporate new neurons in adulthood, in particular the nidopallium. Within this brain region, the boundaries of the song control nucleus HVC could be clearly distinguished from surrounding structures by a higher density of DCX-ir structures. In most telencephalic areas, about two thirds of these cells displayed a uni- or bipolar fusiform morphology suggesting they were migrating neurons. The rest of the DCX-ir cells in the telencephalon were larger and had a round multipolar morphology. No such staining was found in the rest of the brain. The broad expression of DCX specifically in adult brain structures that exhibit the characteristic of active incorporation of new neurons suggests that DCX plays a key role in the migration of new neurons in the brain of adult songbirds as it presumably does during ontogeny.

Six pairs of alipochromatic ('recessive-white') canaries (Serinus canaria) and six pairs of coloured canaries were kept through a complete breeding cycle while being fed a diet providing 12,000 iu vitamin A/kg. The eggs of three pairs (one recessive-white and two coloured) were all unfertilised and there were only 23 hatchlings (14 recessive-white and nine coloured), of which 14 (10 recessive-white and four coloured) were alive after the first moult. However, there was no clinical, biochemical or pathological evidence that the recessive-white canaries were suffering from vitamin A deficiency or that the coloured canaries were suffering from vitamin A toxicity, suggesting that the diet met the vitamin A requirements of both groups.

In this work we analyze the pressure motor patterns used by canaries (Serinus canaria) during song, both in the cases of males and testosterone treated females. We found a qualitative difference between them which was not obvious from the acoustical features of the uttered songs. We also show the diversity of patterns, both for males and females, to be consistent with a recently proposed model for the dynamics of the oscine respiratory system. The model not only allows us to reproduce qualitative features of the different pressure patterns, but also to account for all the diversity of pressure patterns found in females.

The contribution of social factors to seasonal plasticity in singing behavior and forebrain nuclei controlling song, and their interplay with gonadal steroid hormones are still poorly understood. In many songbird species, testosterone (T) enhances singing behavior but elevated plasma T concentrations are not absolutely required for singing to occur. Singing is generally produced either to defend a territory or to attract a mate and it is therefore not surprising that singing rate can be influenced by the sex and behavior of the social partner. We investigated, based on two independent experiments, the effect of the presence of a male or female partner on the rate of song produced by male canaries. In the first experiment, song rate was measured in dyads composed of one male and one female (M-F) or two males (M-M). Birds were implanted with T-filled Silastic capsules or with empty capsules as control. The number of complete song bouts produced by all males was recorded during 240 min on week 1, 2, 4, and 8 after implantation. On the day following each recording session, brains from approximately one-fourth of the birds were collected and the volumes of the song control nuclei HVC and RA were measured. T increased the singing rate and volume of HVC and RA but these effects were affected by the social context. Singing rates were higher in the M-M than in the M-F dyads. Also, in the M-M dyads a dominance-subordination relationship soon became established and dominant males sang at higher rates than subordinates in T-treated but not in control pairs. The differences in song production were not reflected in the size of the song control nuclei: HVC was larger in M-F than in M-M males and within the M-M dyads, no difference in HVC or RA size could be detected between dominant and subordinate males. At the individual level, the song rate with was positively correlated with RA and to a lower degree HVC volume, but this relationship was observed only in M-M dyads, specifically in dominant males. A second experiment, carried out with castrated males that were all treated with T and exposed either to another T-treated castrate or to an estradiol-implanted female, confirmed that song rate was higher in the M-M than in the M-F condition and that HVC volume was larger in heterosexual than in same-sex dyads. The effects of T on singing rate and on the volume of the song control nuclei are thus modulated by the social environment, including the presence/absence of a potential mate and dominance status among males.

Objective: To evaluate the effects of intranasal benzodiazepines (midazolam and diazepam), alpha(2)-agonists (xylazine and detomidine) and their antagonists (flumazenil and yohimbine) in canaries., Study Design: Prospective randomized study., Animals: Twenty-six healthy adult domesticated canaries of both sexes, weighing 18.3 +/- 1.0 g., Methods: In Study 1 an attempt was made to determine the dose of each drug that allowed treated canaries to be laid in dorsal recumbency for at least 5 minutes, i.e. its effective dose. This involved the evaluation of various doses, during which equal volumes of the tested drug were administered slowly into each nostril. In study 2 the onset of action, duration and quality of sedation induced by each drug at its effective dose were evaluated. The efficacy of flumazenil and yohimbine in antagonizing the effects of the sedative drugs was also studied., Results: In study 1 administration of 25 microL per nostril diazepam (5 mg mL(-1) solution) or midazolam (5 mg mL(-1) solution) to each bird caused adequate sedation within 1-2 minutes; birds did not move when placed in dorsal recumbency. After administration of 12 microL per nostril of either xylazine (20 mg mL(-1)) or detomidine (10 mg mL(-1)), birds seemed heavily sedated and assumed sternal recumbency but could not be placed in dorsal recumbency. Higher doses of xylazine (0.5 mg per nostril) or detomidine (0.25 mg per nostril) prolonged sedation but did not produce dorsal recumbency. In study 2 in all treatment groups, onset of action was rapid. Duration of dorsal recumbency was significantly longer (p < 0.05) with diazepam (38.4 +/- 10.5 minutes) than midazolam (17.1 +/- 2.2 minutes). Intranasal flumazenil (2.5 microg per nostril) significantly reduced recumbency time. Duration of sedation was longer with alpha(2)-agonists compared with benzodiazepines. Detomidine had the longest duration of effect (257.5 +/- 1.5 minutes) and midazolam the shortest (36.9 +/- 2.4 minutes). Nasally administered flumazenil significantly reduced the duration of sedation with diazepam and midazolam while yohimbine (120 microg per nostril) effectively antagonized the effects of xylazine and detomidine., Conclusion: Intranasal benzodiazepines produce rapid and effective sedation in canaries. Intranasal alpha(2) agonists produce sedation but not sustained recumbency. Specific antagonists are also effective when used by this route. Clinical relevance Intranasal sedative drug administration is an acceptable alternative method of drug delivery in canaries.

To map the encoding of auditory cues in songbirds, multiunit electrophysiological responses to pure tone stimuli (250-5000 Hz) were recorded at 373 sites throughout the avian analogue of the mammalian auditory cortex in the caudal telencephalon of awake, restrained canaries. We found that a dorso-ventral tonotopic gradient from low to high frequency stimuli extends from the rostral field L2 to caudal-most caudo-medial nidopallium (NCM), similar to the frequency-dependent patterns of ZENK gene expression in canary NCM and to electrophysiological responses in other songbird species. However, response characteristics differ across the region. In field L2, responses are vigorous, phasic, and do not habituate to repeated presentations of the same stimulus. In an important subset of field L2 sites, tuning width narrows over the course of the response, which then terminates rapidly at stimulus offset. These properties are associated with inhibition at many nonpreferred frequencies and poststimulus inhibition at responsive frequencies. In contrast, NCM sites habituate to repeated sine waves, have wider tuning and lower amplitude responses, and rarely show inhibitory effects. Tuning curves in NCM are also flatter than those of field L2, and are often multipeaked. In addition, tuning width increases as the response unfolds and poststimulus excitation is often sustained in NCM. In sum, specific parts of the canary caudo-medial telencephalon differ in their response properties, suggesting differential roles in auditory processing. NCM properties, in particular, may allow for response integration across multiple spectrally varying stimulus elements, such as those that occur during birdsong., (Copyright 2005 Wiley Periodicals, Inc.)

A central aspect of the motor control of birdsong production is the capacity to generate diverse respiratory rhythms, which determine the coarse temporal pattern of song. The neural mechanisms that underlie this diversity of respiratory gestures and the resulting acoustic syllables are largely unknown. We show that the respiratory patterns of the highly complex and variable temporal organization of song in the canary (Serinus canaria) can be generated as solutions of a simple model describing the integration between song control and respiratory centers. This example suggests that subharmonic behavior can play an important role in providing a complex variety of responses with minimal neural substrate.

Masking by harmonic complexes depends on the frequency content of the masker and its phase spectrum. Harmonic complexes created with negative Schroeder phases (component phases decreasing with increasing frequency) produce more masking than those with positive Schroeder phases (increasing phase) in humans, but not in birds. The masking differences in humans have been attributed to interactions between the masker phase spectrum and the phase characteristic of the basilar membrane. In birds, the similarity in masking by positive and negative Schroeder maskers, and reduced masking by cosine-phase maskers (constant phase), suggests a phase characteristic that does not change much along the basilar papilla. To evaluate this possibility, the rate of phase change across masker bandwidth was varied by systematically altering the Schroeder algorithm. Humans and three species of birds detected tones added in phase to a single component of a harmonic complex. As observed in earlier studies, the minimum amount of masking in humans occurred for positive phase gradients. However, minimum masking in birds occurred for a shallow negative phase gradient. These results suggest a cochlear delay in birds that is reduced compared to that found in humans, probably related to the shorter avian basilar epithelia.

Male song complexity is a sexually selected trait found in many songbirds, including strains of the domestic canary. Studies on several species have shown that male song can affect the hormonal state of females and may also influence concentrations of maternal hormones in the yolk of their eggs. In this study, we show that the level of circulating androgens and oestrogens of female canaries, as measured in faeces, varies with the quality of male song to which they are exposed. The female-perceived quality of male canary song depends on the production of special 'sexy syllables' to which females respond with more sexual displays. Using playback of synthetic song we show that females hearing songs with sexy syllables have higher levels of faecal testosterone than control females hearing songs without them. However, unlike previous studies on the canary, we found no evidence that such females laid eggs with more testosterone (or other steroids) in their egg yolks. We discuss these results in relation to the evolution of male signalling and maternal investment strategies.

There is now considerable evidence that female choice drives the evolution of song complexity in many songbird species. However, the underlying basis for such choice remains controversial. The developmental stress hypothesis suggests that early developmental conditions can mediate adult song complexity by perturbing investment in the underlying brain nuclei during their initial growth. Here, we show that adult male canaries (Serinus canaria), infected with malaria (Plasmodium relictum) as juveniles, develop simpler songs as adults compared to uninfected individuals, and exhibit reduced development of the high vocal centre (HVC) song nucleus in the brain. Our results show how developmental stress not only affects the expression of a sexually selected male trait, but also the structure of the underlying song control pathway in the brain, providing a direct link between brain and behaviour. This novel experimental evidence tests both proximate and ultimate reasons for the evolution of complex songs and supports the Hamilton-Zuk hypothesis of parasite-mediated sexual selection. Together, these results propose how developmental costs may help to explain the evolution of honest advertising in the complex songs of birds.

Sexually selected displays, such as male passerine bird song, are predicted to be costly. However, most measurements calculated the rate of oxygen consumption during singing using respirometry have shown that bird song has a low energetic cost. Since birds are reluctant to sing when enclosed inside a respirometry chamber, the energetic cost of singing could differ from that under more normal circumstances. We used heat transfer modelling, based on thermal images, to estimate the energetic cost of singing by canaries (Serinus canaria) that were not enclosed in respirometry chambers. Metabolic rate calculated from heat transfer modelling was 0.70 +/- 0.02 W (N = 10 birds) during singing, which was 14+/-5% greater than during standing (0.62 +/- 0.02 W). The energetic cost of singing did not differ significantly from that measured previously using respirometry when we took into account that birds sang for a greater proportion of the time during the current experiments. These conclusions were not sensitive to potential errors in the heat transfer model. Heat transfer modelling would be especially useful to obtain measurements of the energetic cost of activities that animals do not perform readily inside respirometry chambers, such as singing in birds.

The challenge hypothesis predicts that plasma testosterone (T) concentration is high when male-male competitions are high and decreases when males are engaged in paternal care. In monogamous species, T concentration increases at the beginning of the breeding period and decreases after egg laying. According to the challenge hypothesis, increasing competition should also lead to T increase. The aim of our study was to test this hypothesis. In a first experiment, we measured the T profile of domesticated canaries housed with their mate in separated cages without competition. In a second one, we created a competition by housing male and female domestic canaries together (in an aviary) and emphasized this competition by limiting food access. We also studied social status effect. Our results showed no effect of social status in both sexes and no differences in female's T concentration. Concerning males, we obtained a clear monogamous T profile from the ones housed in a low competition situation and a polygamous profile from the others housed in high competition situation. Thus, our results support the hypothesis of the plasticity of the mechanisms controlling T concentration according to environmental conditions.

Male songbirds such as canaries produce complex learned vocalizations that are used in the context of mate attraction and territory defense. Successful mate attraction or territorial defense requires that a bird be able to recognize individuals based on their vocal performance and identify these songs in a noisy background. In order to learn more about how birds are able to solve this problem, we investigated, with a two-alternative choice procedure, the ability of adult male canaries to discriminate between conspecific song segments from two different birds and to maintain this discrimination when conspecific songs are superimposed with a variety of distractors. The results indicate that male canaries have the ability to discriminate, with a high level of accuracy song segments produced by two different conspecific birds. Song discrimination was partially maintained when the stimuli were masked by auditory distractors, but the accuracy of the discrimination progressively declined as a function of the number of masking distractors. The type of distractor used in the experiments (other conspecific songs or different types of artificial white noise) did not markedly affect the rate of deterioration of the song discrimination. These data indicate that adult male canaries have the perceptual abilities to discriminate and selectively attend to one ongoing sound that occurs simultaneously with one or more other sounds. The administration of a noradrenergic neurotoxin did not impair markedly the discrimination learning abilities although the number of subjects tested was too small to allow any firm conclusion. In these conditions, however, the noradrenergic lesion significantly increased the number failures to respond in the discrimination learning task suggesting a role, in canaries, of the noradrenergic system in some attentional processes underlying song learning and processing.

To what extent seasonal factors modify the neuronal functional properties within the nuclei of the avian song system remains an open question. In adult songbirds, neurons of the song premotor nucleus HVC (used as a proper name) exhibit selective responses for the bird's own song (BOS). Here we examine whether, outside the breeding season, when songs are less stereotyped, HVC neurons of male canaries still respond selectively to the BOS produced during this period. In an initial experiment, single-unit recordings (n = 114) revealed that the neuronal selectivity for the current BOS was attenuated in males exposed to a short-day photoperiod (typical of the nonbreeding season) compared with that found in males exposed to a long-day photoperiod. In long-day conditions, 35% of the cells responded to the BOS, whereas only 12% did in short-day conditions; there were four times more selective cells (d' > 1) in long-day than in short-day conditions. To determine whether these effects were the consequence of differences in acoustic features between breeding and nonbreeding songs, neurons (n = 72) recorded in short-day conditions were tested with both a short-day BOS and a long-day BOS. A low percentage of neurons exhibited responses to short-day or to long-day BOS (11% for each song). Responses of putative interneurons (spike duration < 0.4 ms) and of putative relay cells were similarly attenuated by the short-day conditions. These results strongly suggest that, in canary, rather than being a fixed property, the selectivity for the BOS moves along a continuum and peaks when the day length mimics the breeding conditions.

Canary song is hierarchically structured: Short stereotyped syllables are repeated to form phrases, which in turn are arranged to form songs. This structure occurs even in the songs of young isolates, which suggests that innate rules govern canary song development. However, juveniles that had never heard normal song imitated abnormal synthetic songs with great accuracy, even when the tutor songs lacked phrasing. As the birds matured, imitated songs were reprogrammed to form typical canary phrasing. Thus, imitation and innate song constraints are separate processes that can be segregated in time: freedom in youth, rules in adulthood.

Temperate zone songbirds that breed seasonally exhibit pronounced differences in reproductive behaviors including song inside and outside the breeding season. Springlike long daylengths are associated with increases in plasma testosterone (T) concentrations, as well as with increases in singing and in the volume of several brain nuclei known to control this behavior. The mechanisms whereby T can induce changes in behavior and brain, and whether or not these effects are differentially regulated, have recently begun to be examined, as has the question of the relative contributions of T and its androgenic and estrogenic metabolites to the regulation of this seasonal behavioral and neural plasticity. In this experiment, we examined the effects of T, 5alpha-dihydrotestosterone, or 17beta-estradiol treatment on castrated male canaries housed on short days and compared neural and behavioral effects in these males to similarly-housed males given only blank implants. We observed that only T treatment was effective in eliciting significant increases in singing behavior after 11 days of hormone exposure. In addition, T alone was effective in increasing the volume of a key song production nucleus, HVC. However, at this time, none of the steroids had any effects on the volumes of two other song control nuclei, Area X of the medial striatum and the robust nucleus of the arcopallium (RA), that are efferent targets of HVC, known to be regulated by androgen in canaries and also to play a role in the control of adult song. T can thus enhance singing well before concomitant androgen-induced changes in the song control system are complete.