Behavioral and Brain Sciences Commentary on Keven and Akins --Manuscript Draft-Manuscript Number: Full Title:
Commentary on Keven and Akins
Short Title:
Commentary on Keven and Akins
Article Type:
Commentary Article
Corresponding Author:
Elizabeth Ann Simpson, Ph.D. University of Miami Coral Gables, Florida UNITED STATES
Corresponding Author Secondary Information: Corresponding Author's Institution:
University of Miami
Corresponding Author's Secondary Institution: First Author:
Elizabeth Ann Simpson, Ph.D.
First Author Secondary Information: Order of Authors:
Elizabeth Ann Simpson, Ph.D. Sarah E. Maylott, B.S. Mikael Heimann, Ph.D. Francys Subiaul, Ph.D. Annika Paukner, Ph.D. Stephen J. Suomi, Ph.D. Pier F. Ferrari, Ph.D.
Order of Authors Secondary Information: Abstract:
Empirical studies are incompatible with the proposal that neonatal imitation is arousaldriven or declining with age. Nonhuman primate (NHP) studies reveal a functioning brain mirror system from birth, developmental continuity in imitation and later sociability, and the malleability of neonatal imitation, shaped by the early environment. A narrow focus on arousal effects and reflexes may grossly underestimate neonatal capacities.
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Authors of the target article: Keven and Akins Word counts: Abstract: 59 words; Main text: 837 words; References: 814 words; Entire text: 1,972 words Commentary title: Animal studies help clarify misunderstandings about neonatal imitation Authors of Commentary: Elizabeth A. Simpson, Department of Psychology, University of Miami 5665 Ponce De Leon Boulevard, Coral Gables, Florida 33146, USA. Phone: +1-305-284-6181 Email:
[email protected] Website: http://www.psy.miami.edu/faculty/esimpson Sarah E. Maylott, Department of Psychology, University of Miami 5665 Ponce De Leon Boulevard, Coral Gables, Florida 33146, USA Phone: +1-305-284-1741 Email:
[email protected] Website: http://www.psy.miami.edu/scl/people.phtml Mikael Heimann, Department of Behavioral Sciences and Learning, Linköping University House 1, SE-581 83 Linköping, Sweden Phone: +46-13-281980 Email:
[email protected] Website: http://www.ibl.liu.se/medarbetare/heimann-mikael?l=en&sc=true Francys Subiaul, Department of Speech and Hearing Science, George Washington University 2115 G Street, NW #204, Washington, DC 20052, USA Phone: +1- 202-994-7208 Email:
[email protected] Website: http://www.subiaul.com Annika Paukner, Eunice Kennedy Shriver National Institute of Child Health and Human Development, National Institutes of Health 16701 Elmer School Road, Dickerson, Maryland 20842, USA Phone: +1-301-443-1053 Email:
[email protected] Website: https://www.nichd.nih.gov/about/staff/Pages/bio.aspx?nih_id=0012459403 Stephen J. Suomi, Eunice Kennedy Shriver National Institute of Child Health and Human Development, National Institutes of Health 16701 Elmer School Road, Dickerson, Maryland 20842, USA Phone: +1-301-496-9550 Email:
[email protected] Website: https://www.nichd.nih.gov/about/staff/Pages/bio.aspx?nih_id=0010152932 Pier F. Ferrari, Dipartimento di Neuroscienze, Università di Parma, 43123 Parma, Italy; Institut des Sciences Cognitives Marc Jeannerod, CNRS / Université Claude Bernard Lyon, 67 Bd Pinel, 69675 Bron Cedex, Lyon, France Phone: +39-0521-903947 Email:
[email protected] Website: http://en.unipr.it/ugov/person/17750
Abstract: Empirical studies are incompatible with the proposal that neonatal imitation is arousal-driven or declining with age. Nonhuman primate (NHP) studies reveal a functioning brain mirror system from birth, developmental continuity in imitation and later sociability, and the malleability of neonatal imitation, shaped by the early environment. A narrow focus on arousal effects and reflexes may grossly underestimate neonatal capacities. Main Text: Keven and Akin propose that spontaneous aerodigestive behaviours may be mistaken for neonatal imitation; however, well-designed neonatal imitation studies already account for reflexive and arousaldriven responses (for a review: Simpson et al. 2014a). When measuring arousal, either physiologically or behaviourally, and examining its relationship to imitative responding, evidence shows that for humans (e.g., Nagy et al. 2013) and nonhuman primates (NHP; e.g., Paukner et al. 2016; Simpson et al. 2014b), changes in arousal alone cannot account for neonatal imitation. In addition, Keven and Akin acknowledge that they “have not explained, so far, the differential responses of neonates to specific gestures” (p. 60). We agree and further argue that differential imitation in neonates is incompatible with aerodigestive or arousal-driven explanations. An additional misconception is that neonatal imitation is automatic and involuntary. Instead, infants exert active control over imitative responses and ‘provoke’ previously imitated gestures, even after a delay, in both humans (Meltzoff & Moore 1994) and NHP (Paukner et al. 2011). Moreover, neonates are sensitive both to the type of action and the identity of the individual who modelled the action, initiating interactions only among social partners with whom they previously interacted (Paukner et al. 2011; Simpson et al. 2013). This ability is remarkable because it indicates that newborns are actively socially engaged (Meltzoff & Moore 1994). Consequently, delayed imitation is inconsistent with the proposal that neonatal imitation is a subcortical automatic response. The aerodigestive hypothesis claims that imitative responses peak in the first week of life and decline in the following weeks. The data actually show the opposite for facial gestures: Neonatal imitative responses for tongue protrusion steadily increase in frequency from the first week to the ninth week (e.g., Oostenbroek et al. 2016). Only after 3 months does the frequency of facial gesture imitation decline and infants begin to imitate other actions, such as sounds, vocalizations, and finger movements (Kuhl & Meltzoff 1996; Maratos 1998). Thus, while imitation does undergo changes with development, infants continue to reliably produce matching behaviours (for a review: Simpson et al. 2014a). These findings support the idea that neonatal responses are not stereotypes but rather intentional, voluntary behaviours. We agree with Keven and Akin that animal studies widen our understanding of various phenomena, including neonatal imitation. Nonetheless, in this instance, Keven and Akin neglect to consider animal studies in their potential to inform our understanding of infant social cognitive development (Gerson et al. 2016). For example, Keven and Akin claim, “neonatal imitation experiments provide the only evidence that mirror neurons are present from birth” (pp. 10-11). While we agree that neonatal imitation is behavioural evidence of a functioning mirror neuron system, this assertion overlooks NHP studies documenting cortical brain activity through electroencephalography (EEG) and reporting neural evidence of a mirroring system functioning from birth in newborn monkeys (Ferrari et al. 2012). Furthermore, evidence from animal research is consistent with the premise that individual differences in neonatal imitation may reflect individual differences in sociability (Heimann et al. 1989; Heimann, 1989). Neonatal imitation is hypothesized to be a developmental precursor for, and potentially predict, later social cognitive capacities (Heimann 1991, 2001, 2002; Suddendorf et al. 2013). While this hypothesis has yet to be fully tested in humans, it has been tested in NHP (recent review: Simpson et al. 2016). Infant monkeys who fail to exhibit neonatal imitation, compared to imitators, are less socially attentive (Simpson et al. 2014b), look less at faces in general and the eyes in particular (Paukner et al. 2014), exhibit poorer social cognitive skills such as imitation recognition (Simpson et al. 2015) and gaze following (Simpson et al. 2016), exhibit poorer goal-directed motor skills (Ferrari et al. 2009), play less with peers and exhibit more anxious behaviour at one year of age (Kaburu et al. 2016). Together these
studies provide a more detailed view of the link between neonatal imitation, early social predispositions, and social development. Due to its plasticity, neonatal imitation may also be a fruitful target for intervention, as well as an early marker of sociality. While we know little about the malleability of neonatal imitation in humans (Jacobson 1979; Kennedy-Costantini et al. 2016), animal studies enable the manipulation of infants’ environments and experiences. In monkeys, across both naturalistic and experimental settings, neonatal experiences impact infants’ social capacities (Dettmer et al. 2016), including neonatal imitation, which is strengthened by face-to-face interactions in early infancy (Simpson et al. 2014a; Vanderwert et al. 2015). Contrary to the aerodigestive hypothesis, the aforementioned animal research strongly supports the social nature of neonatal imitation. In sum, evidence to date is inconsistent with the view that neonatal imitation simply reflects spontaneous aerodigestive behaviours. While we appreciate an approach mindful of the broader context of development, there is a wealth of data that directly bear upon Keven and Akin’s arguments. A narrow focus on arousal effects and reflexes may grossly underestimate neonatal capacities. Recognizing such capacities and establishing neonatal measures of sociality may help identify neonates who fall outside the range of healthy social development and may increase opportunities to intervene and foster positive child outcomes. References: Dettmer, A. M., Kaburu, S. S., Simpson, E. A., Paukner, A., Sclafani, V., Byers, K. L., Murphy, A. M., Miller, M., Marquez, N., Miller, G. M., Suomi, S. J. & Ferrari, P. F. (2016) Neonatal face-to-face interactions promote later social behaviour in infant rhesus monkeys. Nature Communications 7:11940. http://www.nature.com/articles/ncomms11940 Ferrari, P. F., Paukner, A., Ruggiero, A., Darcey, L., Unbehagen, S. & Suomi, S. J. (2009) Interindividual differences in neonatal imitation and the development of action chains in rhesus macaques. Child Development 80:1057-1068. http://onlinelibrary.wiley.com/doi/10.1111/j.14678624.2009.01316.x/full Ferrari, P. F., Vanderwert, R. E., Paukner, A., Bower, S., Suomi, S. J. & Fox, N. A. (2012) Distinct EEG amplitude suppression to facial gestures as evidence for a mirror mechanism in newborn monkeys. Journal of Cognitive Neuroscience 24:1165-1172. http://www.mitpressjournals.org/ doi/abs/10.1162/jocn_a_00198#.V73S1ZMrKRs Gerson, S., Simpson, E. A. & Paukner, A. (2016) Drivers of social cognitive development in human and non-human primate infants. In: Frontiers in developmental science series: Social cognition, eds. J. Sommerville, J. Decety. Psychology Press. Heimann, M. (1989) Neonatal imitation, gaze aversion, and mother-infant interaction. Infant Behavior and Development 12:495-505. http://www.sciencedirect.com/science/article/pii/0163638389900295 Heimann, M. (1991) Neonatal imitation: A social and biological phenomenon. In: Behavioral biology: The neuroendocrine axis, ed. T. Archer, S. Hansen, 173-186. Lawrence Erlbaum Associates. Heimann, M. (2002) Notes on individual differences and the assumed elusiveness of neonatal imitation. In: The imitative mind: Development, evolution, and brain bases, eds. A. N. Meltzoff, W. Prinz, 74-84. Cambridge University Press. Heimann, M. (2001) Neonatal imitation: A “fuzzy” phenomenon. In: Emerging cognitive abilities in early infancy, eds. F. Lacerda, C. Von Hofsten, M. Heimann, 231-246. Lawrence Erlbaum Associates. Heimann, M., Nelson, K. E. & Schaller, J. (1989) Neonatal imitation of tongue protrusion and mouth opening: Methodological aspects and evidence of early individual differences. Scandinavian Journal of Psychology 30:90-101. http://onlinelibrary.wiley.com/doi/10.1111/j.14679450.1989.tb01072.x/abstract Kaburu, S. K., Paukner, A., Simpson, E. A., Suomi, S. J. & Ferrari, P. F. (2016) Neonatal imitation predicts infant rhesus macaque (Macaca mulatta) social and anxiety-related behaviours at one year. Scientific Reports. Invited revision submitted. Kennedy-Costantini, S., Slaughter, V. & Nielsen, M. (2016) Why are you copying me? Functional explanations for neonatal imitation. Poster presented at the International Conference on Infant Studies, New Orleans, LA.
Kuhl, P. K. & Meltzoff, A. N. (1996) Infant vocalizations in response to speech: vocal imitation and developmental change. Journal of the Acoustic Society of America 100:2425. http://scitation.aip.org/content/asa/journal/jasa/100/4/10.1121/1.417951 Maratos O. (1998) Neonatal, early and later imitation: same order phenomena? In: The development of sensory, motor and cognitive capacities in early infancy: From perception to cognition, eds. F. Simion, G. Butterworth, 145-160. Psychology Press Ltd. Meltzoff, A. N. & Moore, M. K. (1977) Imitation of facial and manual gestures by human neonates. Science 198:75-78. http://science.sciencemag.org/content/198/4312/75 Meltzoff, A. N. & Moore, M. K. (1994) Imitation, memory, and the representation of persons. Infant Behavior and Development 17:83-99. http://www.sciencedirect.com/science/article/pii/ 0163638394900248 Nagy, E., Pilling, K., Orvos, H. & Molnar, P. (2013) Imitation of tongue protrusion in human neonates: specificity of the response in a large sample. Developmental Psychology 49:1628-1638. http://psycnet.apa.org/journals/dev/49/9/1628/ Oostenbroek, J., Suddendorf, T., Nielsen, M., Redshaw, J., Kennedy-Costantini, S., Davis, J., Clark, S. & Slaughter, V. (2016) Comprehensive longitudinal study challenges the existence of neonatal imitation in humans. Current Biology 26:1334-1338. http://www.sciencedirect.com/science/ article/pii/S0960982216302573 Paukner, A., Ferrari, P. F. & Suomi, S. J. (2011) Delayed imitation of lipsmacking gestures by infant rhesus macaques (Macaca mulatta). PLoS ONE 6:e28848. http://journals.plos.org/plosone/ article?id=10.1371/journal.pone.0028848 Paukner, A., Pederson, E. J. & Simpson, E. A. (2016) Testing the arousal hypothesis of neonatal imitation in infant rhesus macaques. Developmental Psychobiology. Invited revision submitted. Paukner, A., Simpson, E. A., Ferrari, P. F., Mrozek, T. & Suomi, S. J. (2014) Neonatal imitation predicts how infants engage with faces. Developmental Science 17:833-840. http://onlinelibrary.wiley.com/ doi/10.1111/desc.12207/full Simpson, E. A., Murray, L., Paukner, A. & Ferrari, P. F. (2014a) The mirror neuron system as revealed through neonatal imitation: Presence from birth, predictive power, and evidence of plasticity. Philosophical Transactions of the Royal Society B: Biological Sciences 369:1-12. http://rstb.royalsocietypublishing.org/content/369/1644/20130289.short Simpson, E. A., Miller, G. M., Ferrari, P. F., Suomi, S. J. & Paukner, A. (2016) Neonatal imitation and early social experience predict gaze following abilities in infant macaques. Scientific Reports 6:20233. http://www.nature.com/articles/srep20233 Simpson, E. A., Paukner, A., Sclafani, V., Suomi, S. J. & Ferrari, P. F. (2013) Lipsmacking imitation skill in newborn macaques is predictive of social partner discrimination. PLoS ONE 8:1-6. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0082921 Simpson, E. A., Paukner, A., Suomi, S. J. & Ferrari, P. F. (2014b) Visual attention during neonatal imitation in newborn macaque monkeys. Developmental Psychobiology 56:864-870. http://onlinelibrary.wiley.com/doi/10.1002/dev.21146/full Simpson, E. A., Paukner, A., Suomi, S. J. & Ferrari, P. F. (2015) Neonatal imitation and its sensorimotor mechanism. In: New frontiers in mirror neuron research, eds. P.F. Ferrari, G. Rizzolatti, 296-314. Oxford University Press. Suddendorf, T., Oostenbroek, J., Nielsen, M. & Slaughter, V. (2013) Is newborn imitation developmentally homologous to later social‐cognitive skills? Developmental Psychobiology 55:52-58. http://onlinelibrary.wiley.com/doi/10.1002/dev.21005/full Vanderwert, R. E., Simpson, E. A., Paukner, A., Suomi, S. J., Fox, N. A. & Ferrari, P. F. (2015) Early social experience affects neural activity to affiliative facial gestures in newborn nonhuman primates. Developmental Neuroscience 37:243-252. http://www.karger.com/Article/Abstract/ 381538
