A neuroethological view of the multifaceted sensory influences on birdsong.

Learning and execution of complex motor skills are often modulated by sensory feedback and contextual cues arriving across multiple sensory modalities. Vocal motor behaviors, in particular, are primarily influenced by auditory inputs, both during learning and mature vocal production. The importance of auditory input in shaping vocal output has been investigated in several songbird species that acquire their adult song based on auditory exposure to a tutor during development. Recent studies have highlighted the influences of stimuli arriving through other sensory channels in juvenile song learning and in adult song production. Here, we review changes induced by diverse sensory stimuli during the song learning process and the production of adult song, considering the neuroethological significance of sensory channels in different species of songbirds. Additionally, we highlight advances, open questions, and possible future approaches for understanding the neural circuits that enable the multimodal shaping of singing behavior.

Responses in the left and right entopallium are differently affected by light stimulation in embryo.

Sensory stimulation during the prenatal period has been argued to be a main factor in establishing asymmetry in the vertebrate brain. However, though largely studied in behavior and neuroanatomy, nothing is known on the effects of light stimulation in embryo on the activities of single neurons. We performed single-unit recordings from the left and right entopallium of dark- and light-incubated chicks, following ipsi-, contra-, and bilateral visual stimulation. Light incubation increased the general responsiveness of visual neurons in both the left and the right entopallium. Entopallial responses were clearly lateralized in dark-incubated chicks, which showed a general right-hemispheric dominance. This could be suppressed or inverted after light incubation, revealing the presence of both spontaneous and light-dependent asymmetries. These results suggest that asymmetry in single-neuron activity is present at the onset and can be modulated by environmental stimuli such as light exposure in embryos.

Wild nightingales flexibly match whistle pitch in real time.

Interactive vocal communication, similar to a human conversation, requires flexible and real-time changes to vocal output in relation to preceding auditory stimuli. These vocal adjustments are essential to ensuring both the suitable timing and content of the interaction. Precise timing of dyadic vocal exchanges has been investigated in a variety of species, including humans. In contrast, the ability of non-human animals to accurately adjust specific spectral features of vocalization extemporaneously in response to incoming auditory information is less well studied. One spectral feature of acoustic signals is the fundamental frequency, which we perceive as pitch. Many animal species can discriminate between sound frequencies, but real-time detection and reproduction of an arbitrary pitch have only been observed in humans. Here, we show that nightingales in the wild can match the pitch of whistle songs while singing in response to conspecifics or pitch-controlled whistle playbacks. Nightingales matched whistles across their entire pitch production range indicating that they can flexibly tune their vocal output along a wide continuum. Prompt whistle pitch matches were more precise than delayed ones, suggesting the direct mapping of auditory information onto a motor command to achieve online vocal replication of a heard pitch. Although nightingales' songs follow annual cycles of crystallization and deterioration depending on breeding status, the observed pitch-matching behavior is present year-round, suggesting a stable neural circuit independent of seasonal changes in physiology. Our findings represent the first case of non-human instantaneous vocal imitation of pitch, highlighting a promising model for understanding sensorimotor transformation within an interactive context. VIDEO ABSTRACT.

Light-incubation effects on lateralisation of single unit responses in the visual Wulst of domestic chicks.

Since the ground-breaking discovery that in-egg light exposure triggers the emergence of visual lateralisation, domestic chicks became a crucial model for research on the interaction of environmental and genetic influences for brain development. In domestic chick embryos, light exposure induces neuroanatomical asymmetries in the strength of visual projections from the thalamus to the visual Wulst. Consequently, the right visual Wulst receives more bilateral information from the two eyes than the left one. How this impacts visual Wulst's physiology is still unknown. This paper investigates the visual response properties of neurons in the left and right Wulst of dark- and light-incubated chicks, studying the effect of light incubation on bilaterally responsive cells that integrate information from both eyes. We recorded from a large number of visually responsive units, providing the first direct evidence of lateralisation in the neural response properties of units of the visual Wulst. While we confirm that some forms of lateralisation are induced by embryonic light exposure, we found also many cases of light-independent asymmetries. Moreover, we found a strong effect of in-egg light exposure on the general development of the functional properties of units in the two hemispheres. This indicates that the effect of embryonic stimulation goes beyond its contribution to the emergence of some forms of lateralisation, with influences on the maturation of visual units in both hemispheres.