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[eng] This doctoral dissertation examines the effects of a short stay abroad (SA) experience on second language (L2) oral fluency development, taking into consideration individual differences in cognition and the study abroad experience. Relatively little attention has been paid to the cognitive aspects that influence L2 oral fluency gains. Following Segalowitz’s (2010) model, which emphasizes the underlying cognitive processes that affect L2 speech development, the present study investigates both utterance and cognitive fluency, specifically looking at the relationship between inhibitory control (an under-researched aspect of L2 cognition in the SA context) on L2 fluency gains. Forty-nine American students studying Spanish abroad in Barcelona were tested. First, this study investigates whether L2 oral fluency improves over a short stay abroad period, using an in-depth battery of L2 oral fluency measures. Second, we examine the relationship between L2 oral fluency gains and inhibitory control in the SA context, as we hypothesized that greater inhibitory control ability was related to L2 oral fluency gains. Third, we assess the extent to which individual differences in SA experience factors (such as the amount of self-reported language use, living situation and hours of language classes taken) affect L2 oral fluency gains over the SA period. Picture based speech elicitation tasks were used to collect both L1 and L2 oral fluency data, and L2 oral fluency results were adjusted for the L1 where possible. To measure inhibitory control, both non-linguistic and linguistic inhibitory control tasks were used: a Simon task, a Letter (Phoneme) Decision task, and an L1-L2 and L2-L1 language switching task. Participants also completed a post-test questionnaire about their study abroad language use and experience. Results showed little improvement in L2 speed, breakdown, and repair fluency development after a stay abroad. Participants spoke for a longer duration and had longer fluent runs. Contrary to our predictions, after adjusting for the L1, speed fluency data showed that participants spoke significantly slower at the end of their stay, while breakdown fluency results indicate that participants exhibited more silent and filled pauses at the end of their day, only becoming significantly less disfluent on one breakdown fluency measure (between clause silent pauses). A composite disfluency measure showed participants, on average, were less fluent in their L2 at post-test. Inhibitory control ability was not found to relate to L2 oral fluency gains for most measures. A decrease in one disfluency measure –filled pauses between speech units (ASUs) – was significantly related to performance on the Letter (Phoneme) Decision test. Interestingly, participants were able to switch significantly faster into Spanish at the end of their stay than the beginning, which may be one indicator of increased inhibitory control. Most fluency gains were not related to self-reported language use or other experience factors. Only 4 (of 25) fluency measures were significantly related to self-reported language use, one measure was significantly related to classroom instruction hours, and one was significantly related to living situation. The results imply that these specific experiential factors themselves do not have a large impact on participants’ L2 oral fluency gains while abroad, although questionnaire data indicates participants perceived these factors to greatly affect their L2 oral fluency development. Taken together, our findings suggest that a study abroad experience does not necessarily lead to L2 oral fluency gains. Measuring frequency and duration in L2 oral fluency measures and adjusting for the L1 brings insight to the data. Experience factors do not relate to L2 oral fluency gains, contrary to participant expectations and beliefs. Findings from the present study could be applied to pedagogy in the development of future SA programs., [spa] Esta tesis doctoral investiga los efectos de una estancia en el extranjero en el desarrollo de la fluidez de la lengua castellana de un grupo de 49 americanos que estudiaron durante tres meses en Barcelona. Siguiendo el modelo cognitivo de Segalowitz (2010), que se concentra en los procesos cognitivos detrás de la producción oral en segundas lenguas (L2), este estudio investiga tanto el concepto de fluidez en el discurso como la fluidez cognitiva que influye en el discurso. Primero, este estudio evalúa si una estancia corta en el extranjero resulta en una mejora de la fluidez oral, usando una batería de medidas, ajustándolas a la manera individual de hablar en inglés. Segundo, investigamos la relación entre las mejoras en la fluidez y el control de la inhibición, un aspecto de la cognición poco estudiado en lingüística. En tercer lugar, evaluamos cómo las diferencias individuales en su experiencia afectan la mejora del idioma durante la estancia en el extranjero. Los participantes realizaron pruebas del discurso en castellano e inglés, pruebas de inhibición lingüísticas y no lingüísticas, y un cuestionario sobre la experiencia después de la estancia. Los resultados muestran muy poca mejora en la fluidez en el L2 en las tres categorías de fluidez: la rapidez, la disrupción por disfluencias en el discurso y la repetición y autocorrección de errores. Contrario a las expectativas, los participantes hablaban con menos rapidez y fluidez al final de la estancia, en la mayoría de las medidas, menos una – tenían significadamente menos pausas en la mitad de las cláusulas en comparación al principio de la estancia. La inhibición no estaba relacionada con los cambios y mejoras en la fluidez, aunque los participantes pudieran cambiar del inglés al castellano más rápido al final de la estancia. Muy pocas medidas de fluidez estaban relacionadas con los factores de experiencia, indicando que estos factores no influyen en la fluidez, aunque el cuestionario muestra que los participantes lo creen. En conclusión, los resultados de este estudio pueden ser útiles para la pedagogía con el fin de crear nuevos programas de estudio en el extranjero., [cat] Aquesta tesi doctoral investiga els efectes d’una estada a l’estranger en el desenvolupament de la fluïdesa en la llengua castellana d’un grup de 49 americans que van estudiar durant tres mesos a Barcelona. Seguint el model cognitiu de Segalowitz (2010), que es concentra en els processos cognitius darrere de la producció oral en les segones llengües (L2), aquest estudi investiga tan el concepte de fluïdesa en el discurs com la fluïdesa cognitiva que influeix en el discurs. Primerament, aquest estudi avalua si una estada curta a l’estranger resulta en una millora de la fluïdesa oral, utilitzant una bateria de mesures, ajustant-les a la manera individual de parlar l’anglès. En segon lloc, investiguem la relació entre les millores en la fluïdesa i el control de la inhibició, un aspecte de la cognició poc estudiat en la lingüística. Tercerament, avaluem com les diferències individuals en la seva experiència afecten en la millora de l’idioma durant l’estada a l’estranger. Els participants van realitzar proves del discurs en castellà i en anglès, proves d’inhibició lingüística i no lingüística, i un qüestionari sobre l’experiència després de l’estada. Els resultats mostren molt poca millora en la fluïdesa en la L2 en les tres categories de fluïdesa: la rapidesa, la disrupció per disfluències en el discurs i la repetició, i l’autocorrecció d’errors. Contrari a les expectatives, els participants parlen amb menys rapidesa i fluïdesa al final de la seva estada, en la majoria de les mesures, menys una -tenien significativament menys pauses en la meitat de les clàusules en comparació al començament de l’estada. La inhibició no estava relacionada amb els canvis i millores en la fluïdesa, encara que els participants poguessin canviar de l’anglès al castellà més ràpidament al final de l’estada. Molts d’aquests factors no influeixen en la fluïdesa, encara que el qüestionari mostra que els participants ho creuen. En conclusió, els resultats d’aquest estudi poden ser útils per la pedagogia amb la finalitat de crear nous programes d’estudi en l’estranger.

[eng] In my thesis I investigate the evolution of human sociality and its possible role for providing a scaffold to the evolution of language. My approach is multidisciplinary and includes studies ranging from genomic analyses to behavioral experiments in songbirds. My hypothesis is that oxytocin is a good candidate molecule that could help us decipher the role the evolution of our sociality plays in the evolution of language, as well as the role of social reward/motivation in language acquisition. I study the evolution of human sociality mainly through the lens of the ‘self-domestication’ hypothesis, according to which natural selection in humans favored increased prosociality over aggression (Hare 2017), giving rise to a behavioral and an anatomical phenotype that is reminiscent of the one we witness in domesticated species. In Theofanopoulou et al. 2017 (Chapter 1) we examine if this old hypothesis, stemming from thoughts formulated in Darwin 1888, makes sense at a genetic level. We identify a statistically significant intersection of genes under positive selection in modern humans and in one or more domesticated species. Oxytocin/vasopressin receptors are among the genes that have been studied the most in the context of domestication and social cognition. We studied variant changes in these genes between modern humans and our extinct (Neanderthals and Denisovans) and extant relatives (macaques, bonobos, chimpanzees) (Theofanopoulou et al. 2018, Appendix Chapter 1) and report 29 variants which were clustered based on their presence in the species studied and discuss their functional relevance. In Theofanopoulou 2016 (Chapter 2) I put together studies from the literature that point to a role of oxytocin in modulating the multimodality that characterizes our linguistic ability. I follow a bottom up approach, starting off from possible genetic interactions that could support this role, and ending with evidence from EEG (electroencephalography) and behavioral studies. In Theofanopoulou, Boeckx, and Jarvis 2017 (Chapter 3), we propose specific neural mechanisms through which oxytocin could modulate brain regions that are specialized for vocal/speech learning directly, or indirectly through its interaction with dopaminergic neurons. In Theofanopoulou et al. (Appendix Chapter 2) we experimentally address the traditional idea that social reward enhances learning in the realm of vocal learning. In human studies it is not possible to dissociate social reward from vocal learning and study its exact impact on it, so we attempted such a dissociation developing a vocal learning behavioral paradigm with and without social reward in zebra finches. We found that social reward gates their vocal (pitch) learning. In Theofanopoulou et al. (Appendix Chapter 3), we manipulated the oxytocin-system in zebra finches and found that an administration of an oxytocin-antagonist leads to a significant drop in the number of introductory notes in their love song. In Theofanopoulou et al. (Appendix Chapter 4) we study the synteny (genomic territory) of the oxytocin and vasopressin/vasotocin ligands and receptors in 33 vertebrate genomes and 4 invertebrate outgroups and we propose an evolutionary history and a new universal vertebrate nomenclature for all these genes. In conclusion, Ι believe that this thesis offers a fertile ground for future experiments seeking to unravel the effect of social reward in vocal learning developmentally, something that can shed light to the effect that the evolution of our sociality might have had in the evolution of a fully- fledged language in our species. My thesis also lends evidence to a specific hypothesis under which our sociality can be studied, the ‘self-domestication’ hypothesis. Further, the oxytocin and vasotocin systems are shown to be good candidates for uncovering changes that might have had an effect on the evolution of prosociality, but also changes that affect vocal learning behaviors. Lastly, my thesis proposes a universal nomenclature for the vertebrate oxytocin and vasotocin ligands and receptors that is meant to allow easier translation of findings across vertebrates and to foster more informative design of functional experiments across species. References Darwin, C. (1888). The descent of man and selection in relation to sex. Murray. Hare, B. (2017). Survival of the Friendliest: Homo sapiens Evolved via Selection for Prosociality. Annual Review of Psychology, 68(1), 155–186. https://doi.org/10.1146/annurev-psych-010416-044201 Theofanopoulou, C., Andirko, A., & Boeckx, C. (2018). Oxytocin and Vasopressin Receptor variants as a window onto the evolution of human prosociality. BioRxiv, 460584. doi: http://dx.doi.org/10.1101/460584. (Appendix Chapter1) Theofanopoulou, C. (2016). Implications of oxytocin in human linguistic cognition: From genome to phenome. Frontiers in Neuroscience, 10(271). https://doi.org/10.3389/fnins.2016.00271 (Chapter 2) Theofanopoulou, C., Boeckx, C., & Jarvis, E. D. (2017). A hypothesis on a role of oxytocin in the social mechanisms of speech and vocal learning. Proceedings of the Royal Society B: Biological Sciences, 284(1861), 20170988. https://doi.org/10.1098/rspb.2017.0988 (Chapter 3) Theofanopoulou, C., Gastaldon, S., O’Rourke, T., Samuels, B. D., Tiago Martins, P., Delogu, F., … Boeckx, C. (2017). Self-domestication in homo sapiens: Insights from comparative genomics. PLoS ONE 12(10): e0185306. https://doi.org/10.1371/journal.pone.0185306 (Chapter 1) Theofanopoulou, C., Lipkind, D., Tchernichovski, O., Boeckx, C., & Jarvis, E.D. (Appendix Chapter 2). Selective vocal learning in a social reward context. Theofanopoulou, C., Boeckx, C., & Jarvis, E. D. (Appendix Chapter 3). Pilot study: testing the effect of intranasal administration of an oxytocin-receptor antagonist in adult zebra finch directed singing. Theofanopoulou, C., Gedman, G., Cahill, J. A., Boeckx, C., & Jarvis, E.D. (Appendix Chapter 4). A proposed universal nomenclature for the oxytocin and vasotocin ligand and receptor families and their evolutionary history.