Rhythmic entrainment: Why humans want to, fireflies can't help it, pet birds try, and sea lions have to be bribed

Until recently, the literature on rhythmic ability took for granted that only humans are able to synchronize body movements to an external beat-to entrain. This assumption has been undercut by findings of beat-matching in various species of parrots and, more recently, in a sea lion, several species of primates, and possibly horses. This throws open the question of how widespread beat-matching ability is in the animal kingdom. Here we reassess the arguments and evidence for an absence of beat-matching in animals, and conclude that in fact no convincing case against beat-matching in animals has been made. Instead, such evidence as there is suggests that this capacity could be quite widespread. Furthermore, mutual entrainment of oscillations is a general principle of physical systems, both biological and nonbiological, suggesting that entrainment of motor systems by sensory systems may be a default rather than an oddity. The question then becomes, not why a few privileged species are able to beat-match, but why species do not always do so-why they vary in both spontaneous and learned beat-matching. We propose that when entrainment is not driven by fixed, mandatory connections between input and output (as in the case of, e.g., fireflies entraining to each others' flashes), it depends on voluntary control over, and voluntary or learned coupling of, sensory and motor systems, which can paradoxically lead to apparent failures of entrainment. Among the factors that affect whether an animal will entrain are sufficient control over the motor behavior to be entrained, sufficient perceptual sophistication to extract the entraining beat from the overall sensory environment, and the current cognitive state of the animal, including attention and motivation. The extent of entrainment in the animal kingdom potentially has widespread implications, not only for understanding the roots of human dance, but also for understanding the neural and cognitive architectures of animals.