Yoshimasa SEKI

Vocal production learning and beat perception and synchronization (BPS) share some common characteristics, which makes the vocal learning and rhythmic synchronization hypothesis (VLH) a reasonable explanation for the evolution of the capability for rhythmic synchronization. However, even in vocal learners, it is rare to see non-human animals demonstrate BPS to human music. Therefore, the first objective of this article is to propose some possible reasons why we do not see BPS in budgerigars, an excellent vocal learning species, while presenting some of my own findings. The second objective of this article is to propose a seamless bridge to connect the capability for vocal learning and BPS in locomotion. For this purpose, I present my own findings, wherein cockatiels spontaneously sang in synchrony with a melody of human music. This behavior can be considered a vocal version of BPS. Therefore, it can establish a connection between these two capabilities. This article agrees with the possibility that some mechanisms other than the vocal learning system may enable BPS, contrary to the original idea of VLH. Nevertheless, it is still reasonable to connect the capability for vocal learning and that for BPS. At the very least, the capability for vocal learning may contribute to the evolution of BPS. From these arguments, this article also proposes a scenario which includes vocalizing in synchrony as a driving force for the evolution of BPS and the capability for music production.

<title>Abstract</title>Budgerigars (<italic>Melopsittacus undulatus</italic>) are small Australian parrots with a well-documented, learned vocal repertoire and a high degree of vocal production learning. These birds live in large, social flocks and they vocally interact with each other in a dynamic, reciprocal manner. We assume that budgerigars must process and integrate a wide variety of sensory stimuli when selecting appropriate vocal responses to conspecifics during vocal interactions, but the relative contributions of these different stimuli to that process are next to impossible to tease apart in a natural context. Here we show that budgerigars, under operant control, can learn to respond to specific stimuli with a specific vocal response. Budgerigars were trained to produce contact calls to a combination of auditory and visual cues. Birds learned to produce specific contact calls to stimuli that differed either in location (visual or auditory) or quality (visual). Interestingly, the birds could not learn to associate different vocal responses with different auditory stimuli coming from the same location. Surprisingly, this was so even when the auditory stimuli and the responses were the same (i.e., the bird’s own contact call). These results show that even in a highly controlled operant context, acoustic cues alone were not sufficient to support vocal production learning in budgerigars. From a different perspective, these results highlight the significant role that social interaction likely plays in vocal production learning so elegantly shown by Irene Pepperberg’s work in parrots.

We investigated the effects of tapping style on motor performance and neural activity in self-paced and synchronization tapping tasks in three conditions (drum sticking [DS], one-finger tapping [1FT], and four-finger tapping [4FT]). In the synchronization task, participants tapped in synchrony with a metronomic sound. No significant differences were detected in the accuracy of timing control among the tapping styles, whereas larger potentials on EEG waveforms before tap onset were found in 4FT than in DS or 1FT; these may be readiness potentials for the motor commands required to control multiple fingers. As expected, tap intervals were more stable under the synchronization condition than under the selfpaced condition, but no difference was detected in the neural activity evoked before tap onset. Larger neural potentials observed in the early stage after tap onset in DS might be involved in the sensory feedback associated with tool use.

The origin of rhythmic synchronization or entrainment to a musical beat in animals has been widely discussed. Parrots are suitable animals to examine the relationship between the capability of vocal learning and spontaneous rhythmic synchronization. In this study, budgerigars Melopsittacus undulatus learned to tap (peck) 2 keys alternately at a self-paced rate. Then, the metronomic sounds were played in the background during test sessions while the birds were performing the key pecking task, although they were not required to synchronize tap timing with the metronome. We found modest but significant effects of the metronome rhythms on the tap timing in some subjects. We also tested humans Homo sapiens using almost the same method. In contrast to the birds, a number of human subjects synchronized tap timing to the onset of the metronome without verbal or documented instructions. However, we failed to find an effect of the metronome on self-paced tap timing in some human subjects, although they were capable of rhythmic synchronization. This is the first report describing the effects of metronomic sounds on self-paced tapping in nonhuman vocal learners. This study introduces a new method that can be used in future research comparing birds that differ in vocal learning capacities, social structure, age, sex, hormonal status, and so on as part of examinations of the evolutionary foundations of beat processing.

<p>Budgerigars are small parrots and one of the most interesting behaviors in this species is vocal mimicry. This article briefly reviews studies of vocal behavior and the central nervous vocal control system of budgerigars. Other interesting characteristics of this species are behavioral contagion or imitation, and rhythmic synchronization to metronomic stimuli. All of these behaviors are involved in sensory-motor coordination and/or transforming sensory inputs to motor outputs. The neural and psychological substrates of these behaviors may have a link to one another and could give us insight into the biological basis and evolution of vocal learning and communication.</p>