Response facilitation has often been portrayed as a "low level" category of social learning, because the demonstrator's action, which is already in the observer's repertoire, automatically triggers that same action, rather than induces the learning of a new action. One way to rule out response facilitation consists of introducing a delay between the demonstrator's behaviour and the observer's response to let their possible effects wear off. However, this may not rule out "delayed response facilitation" in which the subject could be continuously "mentally rehearsing" the demonstrated actions during the waiting period. We used a do-as-the-other-did paradigm in two orcas to study whether they displayed cognitive control regarding their production of familiar actions by 1) introducing a delay ranging from 60 to 150 sec between observing and producing the actions and 2) interspersing distractor (nontarget) actions performed by the demonstrator and by the subjects during the delay period. These two manipulations were aimed at preventing the mental rehearsal of the observed actions during the delay period. Both orcas copied the model's target actions on command after various delay periods, and crucially, despite the presence of distractor actions. These findings suggest that orcas are capable of selectively retrieving a representation of an observed action to generate a delayed matching response.Moreover, these results lend further support to the proposal that the subjects' performance relied not only on a mental representation of the specific actions that were requested to copy, but also flexibly on the abstract and domain general rule requested by the specific "copy command". Our findings strengthen the view that orcas and other cetaceans are capable of flexible and controlled social learning.

While influential works since the 1970s have widely assumed that imitation is an innate skill in both human and non-human primate neonates, recent empirical studies and meta-analyses have challenged this view, indicating other forms of reward-based learning as relevant factors in the development of social behavior. The visual input translation into matching motor output that underlies imitation abilities instead seems to develop along with social interactions and sensorimotor experience during infancy and childhood. Recently, a new visual stream has been identified in both human and non-human primate brains, updating the dual visual stream model. This third pathway is thought to be specialized for dynamics aspects of social perceptions such as eye-gaze, facial expression and crucially for audio-visual integration of speech. Here, we review empirical studies addressing an understudied but crucial aspect of speech and communication, namely the processing of visual orofacial cues (i.e., the perception of a speaker’s lips and tongue movements) and its integration with vocal auditory cues. Along this review, we offer new insights from our understanding of speech as the product of evolution and development of a rhythmic and multimodal organization of sensorimotor brain networks, supporting volitional motor control of the upper vocal tract and audio-visual voices-faces integration.

Multimodal imitation of actions, gestures and vocal production is a hallmark of the evolution of human communication, as both, vocal learning and visual-gestural imitation, were crucial factors that facilitated the evolution of speech and singing. Comparative evidence has revealed that humans are an odd case in this respect, as the case for multimodal imitation is barely documented in non-human animals. While there is evidence of vocal learning in birds and in mammals like bats, elephants and marine mammals, evidence in both domains, vocal and gestural, exists for two Psittacine birds (budgerigars and grey parrots) and cetaceans only. Moreover, it draws attention to the apparent absence of vocal imitation (with just a few cases reported for vocal fold control in an orangutan and a gorilla and a prolonged development of vocal plasticity in marmosets) and even for imitation of intransitive actions (not object related) in monkeys and apes in the wild. Even after training, the evidence for productive or “true imitation” (copy of a novel behavior, i.e., not pre-existent in the observer’s behavioral repertoire) in both domains is scarce. Here we review the evidence of multimodal imitation in cetaceans, one of the few living mammalian species that have been reported to display multimodal imitative learning besides humans, and their role in sociality, communication and group cultures. We propose that cetacean multimodal imitation was acquired in parallel with the evolution and development of behavioral synchrony and multimodal organization of sensorimotor information, supporting volitional motor control of their vocal system and audio-echoic-visual voices, body posture and movement integration.