Habitat Quality Affects Early Physiology and Subsequent Neuromotor Development of Juvenile Black-Capped Chickadees

In songbirds, the ability to learn and render the species-specific song is influenced by the development of both the song nuclei in the brain and the syrinx (bird''s vocal apparatus) early in the bird''s life. In black-capped chickadees (Poecille atricapillus), habitat quality is known to affect song structure, with birds in high-quality habitat (mature forest) having a higher song consistency than birds in low-quality habitat (young forest). Although this difference is suspected to stem from differences in development, the developmental status of juvenile birds in either habitat remains unexplored. In this study, we used ptilochronology and feather corticosterone to compare the conditional state of juvenile chickadees in young and mature forest during two distinct periods of song learning - the sensory phase, which occurs prior to settlement, and the sensorimotor phase, which occurs post-settlement. A sample of juvenile males was captured and euthanized several weeks prior to their first breeding season to compare the development of song center nuclei and syrinx in both habitats. The corticosterone levels of natally-grown feathers were greater among birds that settled in mature than young forests - as these feathers were grown pre-settlement, they reflect differences in physiology during the sensory phase. This difference in conditional state is reflected by differences in syrinx and song center nuclei development later during the sensorimotor phase - birds in young forest have smaller syrinx, and moderately-larger RA, than birds in mature forest. Those differences could be responsible for the difference in consistency in song structure observed across habitats. The difference in physiological state across habitats, combined with potential compounding effect of differences in winter resources between habitats, could influence the difference in syrinx and neural development seen in juvenile males during the early spring, and influence the male''s ability to learn and render their species-specific song.     

Dynamic model for enzyme action

Background: Protein thermodynamic structure theory is an integrated approach to the study of protein dynamics and the mechanisms of enzyme catalysis. In this paper, a hypothesis arising from this theory is examined. The timescale of an enzymatic reaction (TER) gives a key to characterizing enzyme conformational changes. The aspects of timescale important in our approach are: (i) it is logically related to internal motions of the main chain of a protein; (ii) it sets the upper limit on the size or scope of protein conformational changes. Feature (i) is linked to the dynamic properties of enzyme-reactant complexes. Feature (ii) is linked to the dynamic sites of the main chain (promoting motion) involved in enzyme activity. Conclusion: Our analysis shows that a comprehensive understanding of enzymology can be established on the basis of protein thermodynamic structure theory.     

Processes underlying complex patterns of song trait evolution in a Setophaga hybrid zone

During secondary contact between two species when hybrids are less fit than parents, mating signals are expected to diverge, while aggressive signals are expected to converge. If a single signal trait is used in both mating and aggression, then the dynamics between these two forces could influence the evolutionary trajectory of that trait. We studied such a situation in an avian hybrid zone between two Setophaga species, where birdsong is used in both mate attraction and territory defense. We hypothesized that song modules of the two species will show separate and distinct geographic patterns due to the influence of selective pressures for effective territorial aggression and for effective mate attraction. We conducted geographic cline analyses and playback experiments across this hybrid zone. We found an unexpected geographic pattern of asymmetric introgression of song rhythm, which may be explained by results of the playback experiments that suggest that differences in song rhythm serve a greater role in mate attraction than in territory defense. In contrast, differences in syllable morphology show little evidence of importance in mate attraction or territorial defense. Song features converge in the hybrid zone, yet patterns of trait change suggest that the song production modules may vary in their modes of development and inheritance. Syringeal motor gesturing, which gives rise to syllable morphology, shows a nonclinal mosaic pattern, suggesting that this trait may be predominantly learned. In contrast, respiratory patterning, which forms song rhythm, shows a clinal geographic transition, suggesting that this trait could be more innate. The results indicate that opposing forces act independently on song via distinct modules of the song production mechanism, driving complex patterns of song trait evolution.     

How social experience shapes song representation in the brain of starlings

Birdsong, like speech, is a learned behaviour whose critical function is to communicate with others and whose development critically depends on social influences. Song learning is a complex phenomenon that involves not only the development of species-specific vocalisations, but also the development of the ability to organise these vocalisations and to use them in an appropriate context. Although the fact that interactions with adult experienced models are essential for song production to develop properly has been well established, far less is known about song perception and processing. The fact that songbirds learn to vocalise and to use their vocalisations selectively through interactions with adults questions whether such interactions are also required for songbirds to perceive and process their vocalisations selectively and whether social interactions may shape song perception and processing as they shape song production. In order to address these questions, our team uses an original neuroethological approach to study the neural bases of song behaviour in a highly social songbird species: the European starlings. We provide here a synthesis of the results we have obtained using this approach over the last decade. Our results show that direct social experience with adult experienced models not only shapes song behaviour but also shapes these songbirds’ brains and their ability to perceive and to process acoustic signals whose communicative value, as well as their acoustic structure, have to be learned.     

Limited flexibility in female Pacific field cricket (Teleogryllus oceanicus) exploratory behaviors in response to perceived social environment

How populations adapt, or not, to rapid evolution of sexual signals has important implications for population viability, but is difficult to assess due to the paucity of examples of sexual signals evolving in real time. In Hawaiian populations of the Pacific field cricket (Teleogryllus oceanicus), selection from a deadly parasitoid fly has driven the rapid loss of a male acoustic signal, calling song, that females use to locate and evaluate potential mates. In this newly quiet environment where many males are obligately silent, how do phonotactic females find mates? Previous work has shown that the acoustic rearing environment (presence or absence of male calling song) during late juvenile stages and early adulthood exposes adaptive flexibility in locomotor behaviors of males, as well as mating behaviors in both sexes that helps facilitate the spread of silent (flatwing) males. Here, we tested whether females also show acoustically induced plasticity in walking behaviors using laboratory‐reared populations of T. oceanicus from Kauai (HI; >90% flatwings), Oahu (HI; ~50% flatwings), and Mangaia (Cook Islands; no flatwings or parasitoid fly). Though we predicted that females reared without song exposure would increase walking behaviors to facilitate mate localization when song is rare, we discovered that, unlike males, female T. oceanicus showed relatively little plasticity in exploratory behaviors in response to an acoustic rearing environment. Across all three populations, exposure to male calling song during development did not affect latency to begin walking, distance walked, or general activity of female crickets. However, females reared in the absence of song walked slower and showed a marginally non‐significant tendency to walk for longer durations of time in a novel environment than those reared in the presence of song. Overall, plasticity in female walking behaviors appears unlikely to have facilitated sexual signal loss in this species.     

A method of independent time and frequency decomposition of bioacoustic signals: inter-individual recognition in four species of penguins

The aim of this paper is to introduce a method for analyzing acoustic signals capable of assessing the potential for individual coding information. Signals are analysed both in the time domain (rhythm of emission of the song independent of its frequency content) and in the spectral domain (spectral content of the song independent of the rhythm of emission). The method is then applied to a comparative study of four penguin species, where the problem posed by inter-individual recognition differs from species to species. A direct relationship was shown between the potential of individual coding and the difficulty in panner identification.     

Arrhythmic Song Exposure Increases ZENK Expression in Auditory Cortical Areas and Nucleus Taeniae of the Adult Zebra Finch

Rhythm is important in the production of motor sequences such as speech and song. Deficits in rhythm processing have been implicated in human disorders that affect speech and language processing, including stuttering, autism, and dyslexia. Songbirds provide a tractable model for studying the neural underpinnings of rhythm processing due to parallels with humans in neural structures and vocal learning patterns. In this study, adult zebra finches were exposed to naturally rhythmic conspecific song or arrhythmic song. Immunohistochemistry for the immediate early gene ZENK was used to detect neural activation in response to these two types of stimuli. ZENK was increased in response to arrhythmic song in the auditory association cortex homologs, caudomedial nidopallium (NCM) and caudomedial mesopallium (CMM), and the avian amygdala, nucleus taeniae (Tn). CMM also had greater ZENK labeling in females than males. The increased neural activity in NCM and CMM during perception of arrhythmic stimuli parallels increased activity in the human auditory cortex following exposure to unexpected, or perturbed, auditory stimuli. These auditory areas may be detecting errors in arrhythmic song when comparing it to a stored template of how conspecific song is expected to sound. CMM may also be important for females in evaluating songs of potential mates. In the context of other research in songbirds, we suggest that the increased activity in Tn may be related to the value of song for assessing mate choice and bonding or it may be related to perception of arrhythmic song as aversive.     

Dynamic variation in forebrain estradiol levels during song learning

Estrogens shape brain circuits during development, and the capacity to synthesize estrogens locally has consequences for both sexual differentiation and the acute modulation of circuits during early learning. A recently optimized method to detect and quantify fluctuations in brain estrogens in vivo provides a direct means to explore how brain estrogen production contributes to both differentiation and neuromodulation during development. Here, we use this method to test the hypothesis that neuroestrogens are sexually differentiated as well as dynamically responsive to song tutoring (via passive video/audio playback) during the period of song learning in juvenile zebra finches. Our results show that baseline neuroestradiol levels in the caudal forebrain do not differ between males and females during an early critical masculinization window. Instead, we observe a prominent difference between males and females in baseline neuroestradiol that emerges during the subadult stage as animals approach sexual maturity. Second, we observe that fluctuating neuroestradiol levels during periods of passive song tutoring exhibit a markedly different profile in juveniles as compared to adults. Specifically, neuroestrogens in the caudal forebrain are elevated following (rather than during) tutor song exposure in both juvenile males and females, suggesting an important role for the early consolidation of tutor song memories. These results further reveal a circadian influence on the fluctuations in local neuroestrogens during sensory/cognitive tasks. Taken together, these findings uncover several unexpected features of brain estrogen synthesis in juvenile animals that may have implications for secondary masculinization as well as the consolidation of recent sensory experiences. © 2014 Wiley Periodicals, Inc. Develop Neurobiol 75: 271–286, 2015     

Lesions of a telencephalic nucleus in male zebra finches: Influences on vocal behavior in juveniles and adults

Male zebra finches learn to sing during a restricted phase of juvenile development. Song learning is characterized by the progressive modification of unstable song vocalizations by juvenile birds during development, a process that leads to the production of stereotyped vocal patterns as birds reach adulthood. The medial magnocellular nucleus of the anterior neostriatum (mMAN) is a small cortical region that has been implicated in song behavior based on its neuronal projection to the High Vocal Center (HVC), a nucleus that is critical for adult vocal production and presumably also plays a role in song learning. To assess the function of mMAN in song, ibotenic acid lesions of this brain region were made in juvenile male zebra finches during the period of vocal learning (40-50 days of age) and in adult males that were producing stable song (>90 days of age). Birds lesioned as juveniles produced highly abnormal, poor quality song as adults. Although the overall song quality of birds lesioned as adults was not highly disrupted or abnormal, the postoperative song behavior of these birds was discernibly different due to slight increases in variability of vocal production, particularly at the onset of singing. These results demonstrate that mMAN plays some important role in vocal production during the sensitive period for song learning, and is also important for consistent initiation and stereotyped production of adult song behavior.     

Song learning in zebra finches: some effects of song model availability on what is learnt and when

This study indicates that captive male zebra finches (Taeniopygia guttata) normally learn their song during the juvenile period between independence from their parents and sexual maturity, from whatever suitable song model is available. Virtually nothing is learnt from the father before this time. Hybrid songs may develop if birds are removed from the father and given a new song model before song learning is complete. The absence of a song model during the juvenile stage appears to postpone the sensitive phase and abnormal song is produced until a suitable model becomes available. Normal song will then develop, even in a sexually mature adult. This indicates that experience and not age is the important factor determining the timing of the sensitive phase for song learning in zebra finches.     

Nonstationary Heart Rate Variability in Respiratory Tests

A new method was proposed for processing a nonstationary heart rate by using frequency-modulated signals rather than amplitude-modulated signals equally spaced over several points of time as in the conventional method. A frequency-modulated signal is a set of identical Gaussian peaks that coincide with the true time points of heart beats. A continuous wavelet transform was used to quantitatively describe the heart rhythm signal. A test with controlled breathing was performed as an example and included three consecutive stages: rest, rhythmic breathing at a specified frequency, and exhalation. Tachograms recorded during the breath test was found to be a nonstationary signal with the alternation of peaks of different spectral ranges. A system of quantitative parameters was developed to describe the dynamics of changes in the spectral properties of the tachogram in transitional areas. A static clustering by the effect of the respiratory test and a dynamic clustering in order to identify the time points when the autonomic nervous system is stressed were performed for all subjects. The article discusses the prospects of using the method as a means to analyze the transient effects in various functional tests and as biofeedback that would help to change the heart rhythm.     

Non-Vocal Shaping of Avian Song Development: Parallels to Human Speech Development

Vocal development in young male cowbirds (Molothrus ater) is sensitive to acoustical stimulation from males, but also to social feedback from female cowbirds, even though females do not sing. Juvenile males show different vocal trajectories if housed with local or distant population females. The major goal of the present study was to identify differences in the form and timing of non‐vocal cues from females during the period in early spring when juvenile males begin to sing stereotyped song and to finalize their repertoires. We housed juvenile males with either local or distant population females and no adult males. We found significant differences between the two groups of females in the use of wing stroking and in male reactions to wing strokes and gapes. There were also differences between the groups in male song performance. To understand further the potential consequences of these differences, we correlated measures of male and female responsiveness to results reported in Smith et al. (2000) on vocal ontogeny and song potency. We found that wing stroking by females was associated with a faster rate of song development and tended to relate to differences in song potency. The non‐vocal shaping seen here may represent a general mechanism for the development of vocal communication, as similar processes influence phonological development in human infants.     

脑电信号数据压缩及棘波识别的小波神经网络方法

本文在对小波神经网络及其算法研究的基础上,提出了一种对脑电信号压缩表达和痫样脑电棘波识别的新方法。实验结果显示,小波网络在大量压缩数据的同时,能够较好的恢复原有信号,另外,在脑电信号的时频谱等高线图上,得到了易于自动识别的棘波和棘慢复合波特征,说明此方法在电生理信号处理和时频分析方面有着光明的应用前景。     

Reaction to conspecific degraded song by the wren Troglodytes troglodytes: Territorial response and choice of song post

We investigated the response of the wren Troglodytes troglodytes to playback of a territorial song degraded by long-range propagation. It appears that the wren is sensitive to this degradation since the territorial reaction is less intense with the degraded song than with the undegraded one. However, the degraded song is still considered by the receiver as a specific territorial aggressive signal. This differential response suggests that the male wren can use the degradation characteristics of the signal to adapt its territorial reaction. Indeed, in response to this stimulus, the receiver wren chooses a higher song post. By so doing, the bird improves both the propagation distance of the emitted song and the receiver's ability to hear the opponent's song. This behavioural change may correspond to a communication strategy, counteracting the environmental constraints on sound propagation. Therefore, in response to sound degradation during long-range propagation, birds may have developed behavioural adaptations complementary to the various adaptations concerning song structure and coding-decoding processes.     

A hierarchical neuronal model for generation and online recognition of birdsongs

The neuronal system underlying learning, generation and recognition of song in birds is one of the best-studied systems in the neurosciences. Here, we use these experimental findings to derive a neurobiologically plausible, dynamic, hierarchical model of birdsong generation and transform it into a functional model of birdsong recognition. The generation model consists of neuronal rate models and includes critical anatomical components like the premotor song-control nucleus HVC (proper name), the premotor nucleus RA (robust nucleus of the arcopallium), and a model of the syringeal and respiratory organs. We use Bayesian inference of this dynamical system to derive a possible mechanism for how birds can efficiently and robustly recognize the songs of their conspecifics in an online fashion. Our results indicate that the specific way birdsong is generated enables a listening bird to robustly and rapidly perceive embedded information at multiple time scales of a song. The resulting mechanism can be useful for investigating the functional roles of auditory recognition areas and providing predictions for future birdsong experiments.     

The Steppengrille (Gryllus spec./assimilis): Selective Filters and Signal Mismatch on Two Time Scales

In Europe, several species of crickets are available commercially as pet food. Here we investigated the calling song and phonotactic selectivity for sound patterns on the short and long time scales for one such a cricket, Gryllus spec., available as "Gryllus assimilis", the Steppengrille, originally from Ecuador. The calling song consisted of short chirps (2-3 pulses, carrier frequency: 5.0 kHz) emitted with a pulse period of 30.2 ms and chirp rate of 0.43 per second. Females exhibited high selectivity on both time scales. The preference for pulse period peaked at 33 ms which was higher then the pulse period produced by males. Two consecutive pulses per chirp at the correct pulse period were already sufficient for positive phonotaxis. The preference for the chirp pattern was limited by selectivity for small chirp duty cycles and for chirp periods between 200 ms and 500 ms. The long chirp period of the songs of males was unattractive to females. On both time scales a mismatch between the song signal of the males and the preference of females was observed. The variability of song parameters as quantified by the coefficient of variation was below 50% for all temporal measures. Hence, there was not a strong indication for directional selection on song parameters by females which could account for the observed mismatch. The divergence of the chirp period and female preference may originate from a founder effect, when the Steppengrille was cultured. Alternatively the mismatch was a result of selection pressures exerted by commercial breeders on low singing activity, to satisfy customers with softly singing crickets. In the latter case the prominent divergence between male song and female preference was the result of domestication and may serve as an example of rapid evolution of song traits in acoustic communication systems.