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Decision making resides at the interface between sensory perception and movement production. Female songbirds in the context of mate choice are an excellent system to define neural circuits through which sensory perception influences production of courtship behaviors. Previous experiments by our group and others have implicated secondary auditory brain sites, including the caudal nidopallium (NC), in mediating behavioral indicators of mate choice. Here, we used anterograde tracer molecules to define projections that emerge from NC in female songbirds, identifying pathways through which NC influences downstream sites implicated in signal processing and decision making. Our results reveal that NC sends projections into the arcopallium, including the ventral intermediate arcopallium (AIV). Previous work revealed that AIV also receives input from another auditory area implicated in song preference and mate choice (caudal mesopallium, CM), suggesting that convergent input from multiple auditory areas may play important roles in initiating mate choice behaviors. In the present results, NC projects to an area implicated in postural and locomotory control (dorsal arcopallium, Ad), suggesting that NC may play a role in directing those forms of copulatory behavior. NC projections also systematically avoid a vocal motor region of the arcopallium that is innervated by CM (robust nucleus of the arcopallium, RA). These results suggest a model in which both NC and CM project to arcopallial pathways implicated in behavioral motivation. These brain regions may exert different influences on pathways through which auditory information can direct different facets of behavioral responses to information detected in those auditory signals.

Depression is an umbrella term used to describe a mood disorder with a broad spectrum of symptoms including a persistent feeling of sadness, loss of interest, and deficits in social behavior. Epigenetic research bridges the environmental and genetic landscape and has the potential to exponentially improve our understanding of such a complex disorder. Depression is also a sexually dimorphic disorder and variations exist within epigenetic modification sites between sexes. These sex-specific mediators may impact behavioral symptomology and could serve as therapeutic targets for treatments to improve behavioral deficits. This mini review will focus on the social behavior perspective of depression and specifically explore the sexually different epigenetic modifications on depression.

Research over the last 20 years regarding the link between circadian rhythms and chronic pain pathology has suggested interconnected mechanisms that are not fully understood. Strong evidence for a bidirectional relationship between circadian function and pain has been revealed through inflammatory and immune studies as well as neuropathic ones. However, one limitation of many of these studies is a focus on only a few molecules or cell types, often within only one region of the brain or spinal cord, rather than systems-level interactions. To address this, our review will examine the circadian system as a whole, from the intracellular genetic machinery that controls its timing mechanism to its input and output circuits, and how chronic pain, whether inflammatory or neuropathic, may mediate or be driven by changes in these processes. We will investigate how rhythms of circadian clock gene expression and behavior, immune cells, cytokines, chemokines, intracellular signaling, and glial cells affect and are affected by chronic pain in animal models and human pathologies. We will also discuss key areas in both circadian rhythms and chronic pain that are sexually dimorphic. Understanding the overlapping mechanisms and complex interplay between pain and circadian mediators, the various nuclei they affect, and how they differ between sexes, will be crucial to move forward in developing treatments for chronic pain and for determining how and when they will achieve their maximum efficacy.

The songbird brain contains a network of structures that are specialized for imitative vocal learning. Over the past few years, many new insights have emerged about the structure and function of that system. Among those insights are understanding how specific pathways contribute to specific aspects of vocal behavior, such as control of the acoustic properties of the vocalizations, control of the sequence in which those sounds are produced, and production of calls as well as songs. New research also has indicated that sites outside of the canonical vocal communication system play key roles in the learning, perception, and performance of the sounds used in vocal communication. This chapter details many of those insights and advocates for an expanded perspective on the vocal communication system as a set of interconnected nuclei that are specialized for the production of not only songs but also multiple types of learned signals used in vocal communication. In addition, several new experimental tools and approaches are highlighted that will allow deeper investigations into fundamental questions about the neural basis of learned vocal communication.

Acoustic communication signals are typically generated to influence the behavior of conspecific receivers. In songbirds, for instance, such cues are routinely used by males to influence the behavior of females and rival males. There is remarkable diversity in vocalizations across songbird species, and the mechanisms of vocal production have been studied extensively, yet there has been comparatively little emphasis on how the receiver perceives those signals and uses that information to direct subsequent actions. Here, we emphasize the receiver as an active participant in the communication process. The roles of sender and receiver can alternate between individuals, resulting in an emergent feedback loop that governs the behavior of both. We describe three lines of research that are beginning to reveal the neural mechanisms that underlie the reciprocal exchange of information in communication. These lines of research focus on the perception of the repertoire of songbird vocalizations, evaluation of vocalizations in mate choice, and the coordination of duet singing.

In humans and other animals, behavioural variation in learning has been associated with variation in neural features like morphology and myelination. By contrast, it is essentially unknown whether cognitive performance scales with electrophysiological properties of individual neurons. Birdsong learning offers a rich system to investigate this topic as song acquisition is similar to human language learning. Here, we address the interface between behavioural learning and neurophysiology in a cohort of wild-caught, hand-reared songbirds (swamp sparrows, Melospiza georgiana). We report the discovery in the forebrain HVC of sensorimotor ‘bridge’ neurons that simultaneously and selectively represent two critical learning-related schemas: the bird’s own song, and the specific tutor model from which that song was copied. Furthermore, the prevalence and response properties of bridge neurons correlate with learning ability – males that copied tutor songs more accurately had more bridge neurons. Our results are consistent with the hypothesis that accurate imitative learning depends on a successful bridge, within single cortical neurons, between the representation of learning models and their sensorimotor copies. Whether such bridge neurons are a necessary mechanism for accurate learning or an outcome of learning accuracy is unknown at this stage, but can now be addressed in future developmental studies.

Female songbirds use male song as an indicator of fitness and use that information to select their mate. Investigations of the female auditory system have provided evidence that the neurons within the caudal mesopallium (CM) are involved in the processing of songs that a female finds attractive, however, it is not clear how CM may exert its influence on behavioral indicators of mate choice. In the present study, anterograde tracing revealed the efferent connections of the female songbird CM. The results demonstrate connections to other auditory regions previously described in males, as well as novel connections to brain regions implicated in motor control. As in males, CM neurons in females project robustly to the lateral and medial extents of the caudal nidopallium, and to the ventral intermediate arcopallium. In a novel finding that is not present in males, CM neurons also project to the robust nucleus of the arcopallium and to the caudal striatum. Calling behavior and the expression of copulation solicitation displays are key indicators of female mate choice, and the projections found here bridge critical gaps necessary to understand how auditory perception can influence circuits related to the expression of those affiliative behaviors in female songbirds.

Female songbirds use male song to discriminate among individuals and evaluate their quality as potential mates. Previous behavioral experiments in many species, including the species studied here, have shown that females will solicit copulation in response to song even if no male is present. Those data demonstrate that female mate choice is closely tied to song features, but they leave open the question of which song parameters are most influential in female mate selection. We sought to identify features of male song that are salient for mate choice in female Bengalese finches. Using a novel experimental approach, we simultaneously tested the possible influence of specific notes or note transitions, the number of different note types in the male's repertoire, the complexity of note content and note sequence, and the stereotypy of note content and note sequence. In additional experiments, we also tested the influence of the pitch and tempo of note production. Our results demonstrate that females generally preferred songs containing increased tempo in the context of species-typical frequency bandwidth, consistent with the idea that females prefer songs that are especially challenging to produce. Female preference for song features that pose a neuromuscular challenge has also been reported in other species. Our data extend those observations into a species that thrives in a laboratory setting and is commonly used in studies of the neural basis of behavior. These results provide an excellent new model system in which to study female preference and the neural mechanisms that underlie signal evaluation and mate choice.

A recent study demonstrates how acute neural circuit manipulations can lead to overestimations of circuit function, while chronic manipulations can reveal compensatory modes of plasticity that restore behavior.

Language as a computational cognitive mechanism appears to be unique to the human species. However, there are remarkable behavioral similarities between song learning in songbirds and speech acquisition in human infants that are absent in non-human primates. Here we review important neural parallels between birdsong and speech. In both cases there are separate but continually interacting neural networks that underlie vocal production, sensorimotor learning, and auditory perception and memory. As in the case of human speech, neural activity related to birdsong learning is lateralized, and mirror neurons linking perception and performance may contribute to sensorimotor learning. In songbirds that are learning their songs, there is continual interaction between secondary auditory regions and sensorimotor regions, similar to the interaction between Wernicke's and Broca's areas in human infants acquiring speech and language. Taken together, song learning in birds and speech acquisition in humans may provide useful insights into the evolution and mechanisms of auditory-vocal learning.

Sensorimotor integration is the process through which the nervous system creates a link between motor commands and associated sensory feedback. This process allows for the acquisition and refinement of many behaviors, including learned communication behaviors such as speech and birdsong. Consequently, it is important to understand fundamental mechanisms of sensorimotor integration, and comparative analyses of this process can provide vital insight. Songbirds offer a powerful comparative model system to study how the nervous system links motor and sensory information for learning and control. This is because the acquisition, maintenance, and control of birdsong critically depend on sensory feedback. Furthermore, there is an incredible diversity of song organizations across songbird species, ranging from songs with simple, stereotyped sequences to songs with complex sequencing of vocal gestures, as well as a wide diversity of song repertoire sizes. Despite this diversity, the neural circuitry for song learning, control, and maintenance remains highly similar across species. Here, we highlight the utility of songbirds for the analysis of sensorimotor integration and the insights about mechanisms of sensorimotor integration gained by comparing different songbird species. Key conclusions from this comparative analysis are that variation in song sequence complexity seems to covary with the strength of feedback signals in sensorimotor circuits and that sensorimotor circuits contain distinct representations of elements in the vocal repertoire, possibly enabling evolutionary variation in repertoire sizes. We conclude our review by highlighting important areas of research that could benefit from increased comparative focus, with particular emphasis on the integration of new technologies.

Songbirds are widely used in studies of the neurobiology underlying learning, memory and performance of the sounds used in vocal communication. Development and activity of neurons in many brain sites implicated in those behaviors are closely related to levels of circulating testosterone. Approaches to understand the effects of testosterone in songbirds are presently limited to testosterone implants, which elevate testosterone levels to supraphysiological values, or castration, which eliminates gonadal production of testosterone. Previous studies in mammals indicate that GnRH agonists may be an effective tool to reduce testosterone within that range of extremes and without invasive surgery. To evaluate the effectiveness of the GnRH agonist Deslorelin as a tool to modulate levels of testosterone in songbirds, we recorded the effects of Deslorelin in adult male zebra finches. We recorded songs, body mass and blood testosterone levels pre-treatment, then we gave each bird a small subcutaneous implant of Deslorelin. We measured blood plasma testosterone levels weekly and recorded song behavior and gross morphology of brain, testes and heart at the end of each experiment. Testosterone levels were reduced at the 5mg/kg dose, and the very slight song changes we observed at that dose were like those reported for castrated zebra finches. As expected, there were no changes in the number of cells in androgen-sensitive brain structures. Suppression of testosterone at the 5mg/kg dose was reversible through implant removal. Thus, Deslorelin is a new tool to transiently suppress testosterone levels without the invasiveness and undesirable aftereffects of surgical castration.