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1 Cover Page The handle holds various files of this Leiden University dissertation. Author: Spierings, M.J. Title: The music of language : exploring grammar, prosody and rhythm perception in zebra finches and budgerigars Issue Date:

2 NEDERLANDSE SAMENVATTING Taal is uniek voor mensen. Hoewel alle dieren communiceren, is daarin nog niet dezelfde complexiteit en structuur gevonden als in menselijke taal. Vanwege zijn uniekheid wordt er ook hevig gediscussieerd over het ontstaan en de evolutie van taal. Hoe communiceerden onze verre voorouders? Wat waren de eerste klanken en structuren, en hoe heeft evolutie hierop kunnen doorbouwen? In 1866 liepen de debatten hierover zo op, dat de Société de Linguistique de Paris een ban legde op de discussie over de evolutie van taal. Er was te weinig empirisch bewijs om alle verschillende hypothesen te onderbouwen. Tegenwoordig wordt er al een aantal decennia weer onderzoek gedaan naar de evolutie van taal. Omdat er geen fossielen van gesproken taal bestaan, moet er op een andere manier onderzoek gedaan worden. Een recente, maar belangrijke, manier om empirisch onderzoek te doen naar taalevolutie, is met vergelijkend onderzoek. Hiermee vergelijken onderzoekers de cognitieve eigenschappen van mensen en andere diersoorten. Eigenschappen die gedeeld zijn tussen mensen en andere dieren, zijn de eigenschappen die waarschijnlijk al vroeg in de evolutie ontstaan zijn. Deze eigenschappen kunnen aanwezig zijn geweest in onze verre voorouders en de basis hebben gelegd waarop de evolutie van taal voortgeborduurd kan hebben. Gedurende mijn promotie ben ik onderdeel geweest van een projectgroep waarin we samen vergelijkend onderzoek deden. Deze projectgroep bestond uit wetenschappers die onderzoek deden naar taalontwikkeling bij baby s en wetenschappers die computermodellen maakten voor de betrokken cognitieve processen. Met deze interdisciplinaire groep keken we naar verschillen en overeenkomsten tussen de taalgerelateerde cognitieve eigenschappen jonge mensen en vogels. Daarnaast onderzochten we welke aspecten van taal belangrijk zijn voor mensen en welke eigenschappen computermodellen moeten hebben om het leerproces te kunnen simuleren. In mijn deel van dit project heb ik onderzoek verricht naar cognitieve vaardigheden gerelateerd aan taalperceptie bij twee vogelsoorten, zebravinken (zangvogels) en parkieten (papegaaien). Deze twee dieren staan fylogenetisch ver van de mens af, maar delen één belangrijke eigenschap met mensen: vocaal leren. Zowel zangvogels als papegaaien hebben vocale voorbeelden nodig om zelf hun zang te leren. Dit is vergelijkbaar met hoe mensen taal moeten horen om taal te kunnen produceren. A

3 Addendum Andere primaten, zoals chimpansees en orang-oetangs, zijn geen vocale leerders. Zij produceren dezelfde geluiden, ongeacht of ze een vocaal voorbeeld hebben gehad. Het werk in dit proefschrift bestaat uit twee delen. Ten eerste heb ik onderzoek gedaan naar eigenschappen gerelateerd aan de perceptie van klankstructuren. Hierbij heb ik bijvoorbeeld onderzocht hoe zebravinken lange reeksen geluiden in korte stukken opdelen en of ze gevoelig zijn voor de intonatie van menselijke spraak. In het tweede deel heb ik de complexere eigenschappen onderzocht die gerelateerd zijn aan de perceptie van taal. Hierin heb ik onderzocht of zebravinken en parkieten abstracte structuren kunnen herkennen in geluid, zoals het algemene ritme van een reeks tonen of de grammaticale structuur van een aantal geluidselementen. Perceptie van klankstructuren In hoofdstuk 2 werd onderzocht hoe zebravinken lange reeksen zebravink-geluiden in kortere delen konden opdelen. Ze konden hiervoor gebruik maken van de mate waarin twee klanken na elkaar voorkwamen of, in een tweede groep dieren, van nauwelijks waarneembare stiltes die tussen de delen ingevoegd waren. Mensen kunnen dit allebei vanaf jonge leeftijd al gebruiken om klankreeksen op te delen De resultaten lieten zien dat zebravinken wel deze korte stiltes gebruikten, maar niet welke klanken samen voorkomen. In hoofdstuk 3 werd ook naar lange reeksen gekeken, maar nu van tonen. Deze tonen waren afwisselend hoog-laag in een groep of afwisselend lang-kort in de andere groep. Mensen groeperen dit soort afwisselingen automatisch als groepen met een hoog-laag of kort-lang patroon. De zebravinken groepeerden de afwisselingen in toonhoogte op een vergelijkbare manier met mensen. Ze konden niet leren om onderscheid te maken in tonen die wisselden in toonduur. In hoofdstuk 4 is de perceptie van prosodische patronen onderzocht in zebravinken. Prosodische patronen zijn veranderingen in spraak door het veranderen van de frequentie, duur en amplitude van de lettergreep. Door veranderingen in prosodie kan er een andere betekenis aan dezelfde zin gegeven worden. De zebravinken in deze studie konden goed leren om onderscheid te maken tussen twee verschillende prosodische patronen. Ook als er minder prosodische veranderingen aanwezig waren, maakten ze nog goed onderscheid en zelfs als de spraakklanken veranderden, konden de zebravinken nog het prosodische patroon volgen

4 Nederlandse samenvatting Deze drie studies tonen aan dat de perceptie van prosodische structuren in klanken niet uniek is voor mensen en taal, maar gedeeld is met een zangvogel. Perceptie van abstracte structuren in geluid Voor het begrijpen van taal is het cruciaal dat men de onderliggende structuur van een reeks geluiden kan herkennen. De voorgaande hoofdstukken onderzochten vooral of verschillende klanken onderscheiden en gegroepeerd konden worden. In deze hoofdstukken onderzochten we of vogels de abstracte structuren van een reeks geluiden kunnen leren en generaliseren. In hoofdstuk 5 werd onderzocht of zebravinken en parkieten onderscheid kunnen maken tussen regelmatige en onregelmatige toonreeksen. We testen ze met versnelde en vertraagde versies van de trainingsgeluiden om te zien of ze echt de regelmatigheid hadden geleerd, of dat ze bijvoorbeeld de absolute afstand tussen twee tonen hadden geleerd. De resultaten lieten zien dat beide soorten wel iets van de algemene regelmaat hadden geleerd, maar dat ze ook gelet hadden op details, zoals de precieze lengte van een toon en de lengte van de pauze tussen twee tonen in. Dit onderzoek liet variatie tussen individuen zien, maar toonde ook aan dat generalisatie van abstracte structuren voor zebravinken en parkieten niet vanzelfsprekend is. In hoofdstuk 6 werd dit type onderzoek gerelateerd aan onderzoek naar de structuur in de geluiden die zebravinken produceren. Zowel de zang als andere vocale interacties van zebravinken volgen een bepaalde regelmaat. De combinatie van perceptie en productie maakt zebravinken een goed diersoort voor verder onderzoek naar ritme en regelmaat. Hoofdstuk 7 onderzocht ook of zebravinken en parkieten een abstracte structuren konden herkennen, maar dit keer was het een grammaticale structuur. Ze leerden om onderscheid te maken tussen geluiden in een XYX structuur en geluiden in een XXY structuur. Daarna werden ze getest met nieuwe geluiden die dezelfde structuren volgden. De zebravinken leerden de exacte positie van elk geluid in de training. De parkieten leerden wel de abstracte structuur. Dat betekent dat ze totaal nieuwe geluiden met een XYX structuur in dezelfde categorie plaatsten als de trainingsgeluiden met een XYX structuur. Dit is een van de eerste duidelijk bewijzen dat dieren abstracte structuren kunnen leren en herkennen. A De resultaten van deze en eerdere studies laten zien dat de perceptie van klankstructuren gedeeld is met meerdere soorten. Niet alleen vogels, maar ook ratten

5 Addendum en apen kunnen verschillen en structuren in klanken herkennen. De perceptie van de abstracte structuren van reeksen klanken is daarentegen veel lastiger. Zowel de zebravinken als de parkieten hadden moeite met het leren van het abstracte ritmische structuur van reeksen toontjes. De grammaticale structuren konden wel door de parkieten geleerd worden, maar zij behoren hiermee tot de weinige soorten die dit kunnen. De studies in dit proefschrift laten zien dat vergelijkend onderzoek inzicht kan geven in de cognitieve eigenschappen die aan de basis van taalevolutie kunnen hebben gelegen. Met deze eerste stappen kunnen we verder ontdekken welke onderdelen van taal perceptie wel, en welke niet, aanwezig zijn in verschillende diersoorten. Zo zullen we de volgende puzzelstukjes vinden om steeds meer te ontdekken over het ontstaan van onze taal

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