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The gene encoding the forkhead box transcription factor, FOXP2, is essential for developing the full articulatory power of human language. Mutations of FOXP2 cause developmental verbal dyspraxia (DVD), a speech and language disorder that compromises the fluent production of words and the correct use and comprehension of grammar. FOXP2 patients have structural and functional abnormalities in the striatum of the basal ganglia, which also express high levels of FOXP2. Since human speech and learned vocalizations in songbirds bear behavioral and neural parallels, songbirds provide a genuine model for investigating the basic principles of speech and its pathologies. In zebra finch Area X, a basal ganglia structure necessary for song learning, FoxP2 expression increases during the time when song learning occurs. Here, we used lentivirus-mediated RNA interference (RNAi) to reduce FoxP2 levels in Area X during song development. Knockdown of FoxP2 resulted in an incomplete and inaccurate imitation of tutor song. Inaccurate vocal imitation was already evident early during song ontogeny and persisted into adulthood. The acoustic structure and the duration of adult song syllables were abnormally variable, similar to word production in children with DVD. Our findings provide the first example of a functional gene analysis in songbirds and suggest that normal auditory-guided vocal motor learning requires FoxP2.
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Vocal imitation plays a fundamental role in human language acquisition from infancy. Little is known, however, about how infants imitate other's sounds. We focused on three factors: (a) whether infants receive information from upright faces, (b) the infant's observation of the speaker's mouth and (c) the speaker directing their gaze towards the infant. We recorded the eye movements of 6-month-olds who participated in experiments watching videos of a speaker producing vowel sounds. We found that an infants' tendency to vocally imitate such videos increased as a function of (a) seeing upright rather than inverted faces, (b) their increased looking towards the speaker's mouth and (c) whether the speaker directed their gaze towards, rather than away from infants. These latter findings are consistent with theories of motor resonance and natural pedagogy respectively. New light has been shed on the cues and underlying mechanisms linking infant speech perception and production.
Adult caregivers display several vocalization topographies that may facilitate social responses and prompt infant vocal responses during face-to-face interactions (e.g., describing ongoing events, imitating, naming objects). Among these forms, maternal imitation of infant vocal sounds has been scarcely studied. Maternal imitation of infant vocalizations is a form of stimulation similar in topography to the responses produced by the child. A number of studies have shown that parents naturally imitate various acoustic features produced by their infants (Field, 1977; Papousek, 1992; Pawlby, 1977). Masur (1987) reported that mothers were likely to follow their infants' vocal imitations with a repetition of the imitation. Masur and Olson (2008) analyzed naturally occurring maternal vocal topographies in response to infant vocalizations during routine caretaking tasks (free play and bathtime) in twenty 10-month-old infants and their mothers. These authors reported that mothers responded to infant vocalizations on 86% of the opportunities and that maternal imitations occurred on 21% of these opportunities.
A small body of research suggests the influence of maternal imitation on the behavior of infants. Field, Guy, and Umbel (1985) found that 3.5-month-old infants vocalized and smiled more frequently subsequent to maternal imitative as opposed to nonimitative behavior during both spontaneous and imitative face-to-face interactions. Masur and Olson (2008) showed that infant behavior was highly responsive to maternal imitations (e.g., 90% of opportunities in 10-month-old infants) whether in the form of return vocal imitation or other social responses (e.g., smiles, gaze shifts, object-related behaviors, nonimitative vocalizations). Responsiveness to maternal imitation, in terms of the proportion of occasions on which children used words to respond to maternal imitation, was correlated with children's lexicon at 21 months of age (see also Tamis-LeMonda, Bornstein, & Baumwell, 2001). Gadzag and Warren (2000) investigated the effects of adult contingent vocal imitation on acquisition of vocal imitation in three young children with mental retardation and showed increased spontaneous imitation during generalization sessions. Based on this literature, we sought to investigate the effects of contingent maternal imitation of infant vocal sounds on vocalizations of infants 3 to 8 months of age.
Bloom (1975) compared the rate of vocalization in 3-month-old infants across phases of contingent and noncontingent adult social stimulation. Both interventions increased rates of infant vocalizations. The author concluded that infant vocalizations may be elicited, rather than reinforced, by adult social stimulation (see Bloom, 1979, for a discussion). However, her research could not rule out the potential for adventitious reinforcement of infant vocalizations during the NCR control condition. Unlike NCR, DRO schedules reduce the possibility of adventitious or intermittent reinforcement of infant vocalizations.
Although the reinforcing effect of other forms of parental stimulation, including talking, touching, and presenting leisure items, has been demonstrated using adequate reinforcement control conditions (e.g., Poulson, 1983, 1988), the effect of maternal imitation as a reinforcer for infant vocalization has not been explored systematically. Furthermore, previous studies that used maternal vocalization as a reinforcing stimulus did not control for specific forms of adult response to infant vocalizations (e.g., Poulson, 1983). In addition, neither NCR nor DRO controls have been used to test the reinforcing effect of maternal vocal imitation on infants' early vocal sounds.
To ensure that all mothers correctly imitated their infants' vocalizations during CR1 and CR2, and also to ensure even distribution of reinforcer density across CR1 and DRO, we monitored and coded all maternal vocal responses. The first author discontinued the session in the event that the mother did not imitate an instance of infant vocalization correctly. Specifically, if the latency to maternal imitation was more than 2 s or the sequence of sounds by the mother (vowel and consonants) did not match that of the infant, the principal investigator discontinued the session. Research assistants retrained mothers as needed before resuming the protocol. In addition, the dyad was excluded if the mother's performance was inconsistent across CR1 and DRO more than five times as a result of not following instructions.
Future studies would benefit from longer sessions, more sessions within phases, and a second DRO control condition in which mothers' vocalization frequency could be yoked to the second CR condition (Thompson & Iwata, 2005). However, the current results suggest that contingent maternal vocal imitation reinforces infant vocalizations within a few treatment sessions.
Previous experiments have only tested the signal function of imitation within dyadic interactions with playback to a single subject at a time [16]. In this experiment, we create a simple communication network by joining two wild-caught orange-fronted conures from different flocks in the same aviary and presenting playback stimuli to them simultaneously. Our aim was to determine if specific receivers can be addressed in a non-territorial communication network. To do so, we simulated vocal imitation through playback of variants of contact calls similar to that of an intended receiver. If vocal imitation addresses specific individuals, we expected the imitated individual to be the primary respondent to the playback. Orange-fronted conures respond with most calls in interactions with the opposite sex and where female test-birds generally gave more calls than male test birds. Given the strong influence of sex in other experiments [16], we expected an overall stronger response of female than male test birds. Most experiments on vocal matching have only monitored the vocal response of the focal bird [4], [5], [8], [20] although other individuals of the local network respond after eavesdropping on the experiment [21],[22],[23],[24]. Our experimental setup, however, enabled us to monitor the vocal response of the whole communication network rather than just the focal bird.
On basis of the audio recordings of each trial, we extracted and logged the time of every contact call given from the two birds in the aviary. To identify the vocalising individuals, we examined the video recordings of the experiment.
Our experiment monitored the response of both intended and non-intended receivers to a sender that did not vary its calls during the interaction. In some natural interactions, the degree of call modification is asymmetric among the interactants; immitation often results from one individual modifying its call more the other individual, rather than a mutual convergence. The playback therefore laid within the natural range of behaviours in orange-fronted conures. Natural interactions between members of flocks would not allow us to discern who addressed who as both flocks (i.e. communication networks) may attempt to address each other. Only an experiment would allow such conclusions. The receiver responses, therefore, are essential for testing whether imitations of contact calls can address other individuals and, ultimately, for understanding the function and evolution of vocal imitation. be457b7860
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