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LINGUIST List 25.2334

Tue May 27 2014

Review: Cognitive Science, General Linguistics: Bolhuis and Everaert (Eds., 2013)

Editor for this issue: Anja Wanner <anjalinguistlist.org>

Date: 21-Nov-2013
From: Darcy Sperlich <darcy.sperlichmanukau.ac.nz>
Subject: Birdsong, Speech, and Language
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Book announced at http://linguistlist.org/issues/24/24-1468.html

EDITOR: Johan J. Bolhuis
EDITOR: Martin Everaert
TITLE: Birdsong, Speech, And Language
SUBTITLE: Exploring the Evolution of Mind and Brain
YEAR: 2013

REVIEWER: Darcy Sperlich, Manukau Institute of Technology


This books revolves around the theme of what the study of birdsong can contribute to our understanding of human language evolution. This being a very broad topic, the book is divided into six comprehensive sections, ‘Introduction,’ ‘Acquisition of birdsong and speech,’ ‘Phonology and syntax,’ ‘Neurobiology of song and speech,’ ‘Genes, song, speech and language,’ and finally ‘Evolution of song speech and language.’ The book begins with a foreword from Robert C. Berwick and Noam Chomsky, setting the scene with Aristotle’s observations on birdsong. They discuss evolutionary problems along the lines of shared ancestry versus functional adaptations, and how this provides insights to the origins of language. This is then followed by the editors’ preface to the book.

The first chapter of the book is entitled ‘The design principles of natural language,’ by Martin Everaert and Riny Huybregts, which is written as an introduction to non-linguists on the principles behind the book. They give a brief overview of what language and generative grammar is and cover perspectives on the design principles behind language. The discussion then turns to that of birdsong, covering arguments that birdsong has grammar on par with humans. They conclude briefly on the evolution of language.

The next chapter entitled ‘Evolution, memory and the nature of syntactic representation,’ by Gary F. Marcus, focuses on the psychological reality of syntactic tree structure in the brain. He proposes it does exist, linked via small branches (treelets), not as a single tree. He divulges into the psychological aspects (e.g., problems of center-embedding), arguing for the treelet approach.

The third article, ‘Convergence and deep homology in the evolution of spoken language,’ by W. Tecumseh Fitch and Daniel Mietchen focuses on the genetic aspects of language. They set the background by discussing homology in evolutionary biology, and then turn to ‘deep homology’ whereby the genetics over different species appear to be similar to one another. One example give is the FOXP2 gene, which is similar to genes found in mice and other vertebrates. They point out how such similarities should not be surprising, and how research on one species can inform on another.

Chapter 4 by Erich D. Jarvis, entitled ‘Evolution of brain pathways for vocal learning in birds and humans,’ focuses on vocal learning. He points out that while most vertebrates are capable of auditory learning, the same is not true of vocal learning. He then compares various brain structures, and how legions of the brain affect vocal behaviour. This is followed by studies on brain activity and genes. The focus is then shifted to the auditory system, looking at the evolution of humans and birds from an earlier ancestor, proposing a motor theory for vocal learning, advancing on earlier theories (e.g., Lieberman 2002).

The next book section, ‘Acquisition of birdsong and speech,’ starts with Sanne Moorman and Johan J. Bolhuis’ comparisons of just this in their short article ‘Behavioral similarities between birdsong and spoken language.’ They go over the basis of avian vocalisation, drawing parallels between how birdsong is learnt compared to language. They discuss sensitive learning periods in birds akin to the critical period hypothesis compared to humans, and finally, syntactic parallels, which they call into question.

‘Parametric Variation: Language and Birdsong,’ by Neil Smith and Ann Law, looks at what makes human language unique, compared to birds. They briefly consider recursion and reject it in favour of parametric variation (PV, of Principles and Parameters theory), which is basically parameter setting (e.g., plus or minus prodrop). After outlining the theory, they list 7 criteria of PV. They then move to birds, comparing and contrasting the crucial differences humans, in order to see if birds meet the PV criterion. They find that, while PV is helpful as a tool, they are unable to conclude if PV is unique to language.

The next article by Olga Fehér and Ofer Tchernichovski, called ‘Vocal culture in songbirds: an experimental approach to cultural evolution,’ is about vocal cultural evolution in birds. It is an experimental study where they isolate zebra finches from the wild and put them in an isolated colony. Once the isolated bird has developed its own song, they then have it ‘tutor’ a younger bird to see what is learnt, repeating the process with the younger bird to other younger birds (also to a group as well). They find over a few generations the song progresses towards to what is found in the wild (the analysis is discussed in detail). They conclude by discussing likely models of this evolution.

Chapter 8 by Frank Wijnen, ‘Acquisition of linguistic categories: cross-domain convergences,’ explores the learning processes involved in syntax and phonology, arguing that they are essentially the same using linguistic definitions as evidence. He covers learning of phonemes and grammatical categories in children, proposing the same statistical learning mechanism underlies their acquisition, being able to track the quantity of tokens and their dependencies in a series. He then reviews evidence in support of this in neuropsychological studies, noting that motor skill acquisition is linked with non-linguistic sequence learning. He concludes by showing how his research paradigm differs from theories of similar nature.

The next chapter, ‘Structure in human phonology and in birdsong: a phonologist’s perspective,’ by Moria Yip, starts the next section called ‘Phonology and syntax.’ She begins by comparing sounds produced by birds and humans in terms of phonological structure. This is done over many species of birds, investigating acoustic, neurological and behavioural cues. Yip then investigates how structure arose in birds (e.g., genetic mutations), exploring five possibilities. She sums up by suggesting further areas to gather evidence.

Eric Reuland in ‘Recursivity of language: what can birds tell us about it?’ focuses on the centrality of recursion being to human language (e.g., Hauser et al., 2002). This is discussed with reference to Minimalism (e.g., Merge), outlining the human computational system. He discusses what language needs, and argues for recursion being a key component. He finally refers to recent work on the European starling, arguing that it does not have recursion. He comes to the conclusion that studying birds will shed little light on human language recursion.

From the outset, Kazuo Okanoya in ‘Finite-state syntax in Bengalese finches: sensorimotor evidence, developmental processes, and formal procedures for syntax extraction’ forcefully argues that birds do not have syntax on par to humans, but rather a finite state syntax. Okanoya moves onto the discussion of segmentation and chunking, asking if birds are able to perceive and produce in such a manner. Reviewing experiments across different birds supported just this, implying a possible cognitive basis.

In a review article entitled ‘Analyzing the structure of bird vocalization and language: finding common ground,’ Carel ten Cate, Robert Lachlan and Willem Zuidema discuss the methods used to identify the structure of songs in birds, and the difficulties posed doing this across species (also in humans). The authors then cover the sequence of song and its complexity – the models used to describe it with special reference to hidden Markov models.

Chapter 13, ‘Phonological awareness in grey parrots: creation of new labels from existing vocalizations,’ by Irene M. Pepperberg, focuses on the famous grey parrot Alex (now deceased), and his colleague, Arthur. The discussion is of the human-like phonological ability they have, due to training and simple human language exposure. The article details the research that has gone into the two parrots, noting the developmental difference between them owing to their differing experimental treatment.

‘The neural basis of links and dissociations between speech perception and production’, the 14th contribution, by Sophie K. Scott, Carolyn McGettigan and Frank Eisner, begins the 4th section of the book, ‘Neurobiology of song and speech.’ They start by discussing how speech perception is represented in the brain, followed by its production, and their relation to the motor cortex. They finally give a summary discussion on the processes, arguing against motor theory of speech perception.

Sharon M. H. Gobes, Jonathan B. Fritz and Johan J. Bolhuis look into auditory learning similarities in songbirds and speech learning in humans, as well as other non-learning species. They briefly cover the literature, then move into auditory learning in nonhuman species, comparing the avian and mammalian brain. The authors then discuss song locality in the bird’s brains and also the neural makes of song memory. They also discuss auditory memory in humans, finishing the chapter with the interaction between auditory and vocal regions, touching on evolutionary matters.

In the 16th chapter of the volume, ‘Age effects in language acquisition and attrition’, Christophe Pallier investigates the issues of language acquisition from an age perspective, tackling the notion of the critical period hypothesis. The review shows that the idea of the brain losing its plasticity is a simplistic approach. Through reviewing the delay of L1 and L2 acquisition, to interesting studies on adopted children growing up in a completely different L1 environment, points to a complex matrix yet to be fully understood.

The next paper, ‘A “birdsong perspective” on human speech production,’ by Hermann Ackermann and Wolfram Ziegler, compares the avian song within the brain to human speech production. They discuss first the similarities between both (e.g., function) and discuss the area of the brain connected with song production, similarly done for humans as well (bringing in studies on brain damage).

Michale S. Fee and Michael A. Long in ‘Neural mechanisms underlying the generation of birdsong: a modular sequential behavior’ investigate the neurology behind song, focusing specifically on the interaction between RA and HVC neurons. After reviewing their functions, the authors move on to two models of song generation, which make use of these neurons, discussing how to experimentally investigate the models and detailing such past experiments by the authors. They finally consider models of timing syllable and motif onsets, and compare them to the data.

The article ‘Auditory-vocal mirror neurons for learned vocal communication,’ by Jonathan F. Prather and Richard Mooney, focuses on mirror neurons in birds. They first discuss the advantages for looking for mirror neurons in songbirds, followed by a discussion on songs in terms of neurons involved and structure in the brain. They then move into experimental evidence measuring neuron activity. The most important point is that HVCX cells appear to be mirror neurons. The authors finally discuss HVCX neurons place in perception, learning and sensorimotor learning, concluding on a mechanism for auditory-vocal correspondence.

Chapter 20 entitled ‘Peripheral mechanisms of vocalization in birds: a comparison with human speech,’ by Gabriël J. L. Beckers, focuses on reviewing vocal production by birds and comparing them to humans. The author discusses the physical mechanisms behind vocal production in birds, e.g., the syrinx, which is involved in the production and modulation of voice. The voice tract’s production use follows, and finally suggests future research directions.

Chapter 21 introduces the next section, ‘Genes, song, speech and language,’ with a paper titled ‘Building bridges between genes, brains and language,’ by Simon G. Fisher. He reviews FOXP2 (in italics), focusing on what it is and what it is not. He covers the discussion of FOXP2 and its links between it and speech – noting that it cannot be seen as a ‘gene for speech’. He covers FOXP2’s functions and areas it affects within the brain, also observing mutations of a similar gene in mice. He finishes his article by looking at the gene’s role in speech evolution.

Constance Scharff and Christopher K. Thompson’s contribution, ‘A bird’s-eye view of FoxP2,’ relates FoxP2 in Area X. They discuss the gene in pre- and postnatal development, within seasonal changes (in canaries) and in general behaviour. The authors touch upon a variety of other topics relating to FoxP2, finishing by looking at the future of genetics and birds.

Chapter 23, titled ‘Genetic basis of language: insights from developmental dyslexia,’ by Franck Ramus, discusses what developmental dyslexia can show us about genes related to language. He backgrounds dyslexia and its main symptoms. He then moves onto the review of gene studies related to the topic, where evidence points to genetic problems causing dyslexia. A similar discussion is given of specific language disorder, and how it overlaps with dyslexia.

W. Tecumseh Fitch leads the final section of the book, ‘Evolution of song, speech and language,’ with ‘Musical protolanguage: Darwin’s theory of language evolution revisited.’ The topic is musical protolanguage (a term coined by Fitch, but a theory of Darwin’s). He reviews the hypothesis and draws parallels to today’s linguistic endeavours. Not only does he discuss its good points, but also its difficulties (clashing with modern linguistics, sexual selection and his terminology), with Fitch suggesting solutions to these problems.

Chapter 25, ‘Birdsong as a model of studying factors and mechanisms affecting signal evolution,’ by Kazuo Okanoya, provides a different perspective on birdsong, focusing on the difference between the Bengalese finch and white-rumped munia, which share a close genetic relationship. He covers a variety of interesting studies which show how different the two species are in their song abilities. He concludes on the developmental pathway of birdsong, and calls for further research.

The final chapter, by Irene M. Pepperberg, ‘Evolution of vocal communication: an avian model,’ focuses on mirror neurons and their role played in evolution, touting birds as a possible missing link in regard to vocal communication. She covers language evolutionary theory and discusses the role of mirror neurons, paralleling bird and human similarities – all the while looking at what species could bridge the divide between the two.


The first chapter by Everaert and Huybregts sets the mood of the book, but it should be noted that the following chapters are not necessarily in concordance of their views espoused here. In saying that, however, they do give consideration to competing thoughts. On another note, I was partly surprised to see the Wikipedia reference on language in the text, which shows how far this encyclopedia has come. In all, serving as an introduction for non-linguists, the authors do the job well.

The article by Marcus presents an attractive idea on treelets, offering courses of research that could help forward the idea. My only comment on this idea would be its appeal to ‘sequence sensitive structure’ present (e.g., the active and passive), but I wonder how useful this paradigm would be for free word order languages, as those found in Australian aboriginal languages (e.g., Jiwarli; Austin 2001). Also, another area would be that of reflexives, which a syntactic approach is concerned with locality, but may run into difficulty trying to explain long distance dependencies (e.g., long distance anaphora, cf. Huang 2000).

The article by Fitch and Mietchen gives an excellent, accessible overview into why we should be studying deep homology across different species, painting a bright future for this field. I found the discussion on the genes involved quite clear, and it also acts as a warm up to the gene related studies presented later on in the volume.

Chapter 4 by Jarvis gives a thorough look at vocal learning from the brain and genes, which is full of references. The topics are well discussed, and the reader comes away with up-to-date knowledge on the latest developments on comparisons between birds and humans. This chapter again is quite useful pre-reading for later chapters.

Moorman and Bolhuis in Chapter 5 is straight to the point in describing parallels between birds and humans, including syntactic parallels which is more controversial, the authors acknowledge (as other articles in the volume also discuss). If there were syntactic parallels, then this obviously would end at a very early stage, otherwise we would expect bird language.

Smith and Law try to answer the question what makes human language unique by introducing parametric variation. In the end they are unable to conclude if parametric variation is unique to language. While it appears useful as a tool of language classification, I would share the authors’ muted conclusion surrounding the uniqueness of parametric variation to language, as to me it would seem not to be the case.

The paper by Fehér and Tchernichovski discusses how birdsong can become wild-like from an experimental setting over a few generations, using objective measures. Cleary, nature and nurture both have a place in these results.

Wijnen’s look into finding an underlying mechanism into phonological and syntactic processes appears very promising on the evidence provided. Such a proposal would find opponents (e.g., Wexler 2011) who argue against statistical learning. Nonetheless, Wijnen’s work adds more to the debate.

Yip’s article looks at the kinds of structure in birdsong over many different species, so in a sense it is more like a meta-analysis. While the search for structure appears hopeful, this would need to be investigated deeply in individual species. Overall, Yip provides a good perspective on the issues at hand, and explains concepts clearly to the non-phonologists.

The article by Reuland presents a Minimalist solution to the question of the uniqueness of human language and sums up the perspective well. Moreover, he does not warrant research into birdsong for recursion as he believes that little will be found -- which may be the case.

Okanoya presents an interesting study on bird syntax through experiments on perception and production, making a case for birds’ lack of recursive syntax, instead having a finite state grammar. Undoubtedly, such research on a single species gives the necessary depth on the status of their syntax, and paints a bright picture for further research in this area.

Cate et al. provides a useful review of the literature on the structure of birdsong, and offers productive steps forward towards a systematic study of birdsong compared to human language.

Pepperberg’s article gives an interesting insight on the phonological acquisition of Alex, which draws parallels to human phonological acquisition. While Alex has died, research continues to be published on his abilities (e.g., on his mathematical ability, Pepperberg 2012). Hopefully, other grey parrots will be able to be similarly raised to provide collaborating evidence.

Scott et al. provide a methodological discussion on the literature of speech perception and production, pointing out that their supposed similarities of processing is not reflected by current evidence. The authors make this point convincingly, for as usual the picture is often more complicated than earlier thought -- and it is good that we are digging deeper into the problem.

Gobes et al. give a state of the art review of the auditory literature, comparing and contrasting the avian and mammalian brains. In all, the review is written well imparting an objective view on the topic.

Pallier’s contribution brings together L1 and L2 studies on language acquisition, effectively showing the problems surrounding the critical period hypothesis. I was especially interested in the research on bringing up adopted children in a new L1 environment, and how much of their L1 they had lost at a young age.

Ackermann and Ziegler’s contribution gives a clear picture on the neurological basis of song in birds and human speech production. Each area of discussion is written concisely, allowing the reader to have a clear understanding of the processes behind these similar systems.

Coming from a linguistic perspective, I found fascinating Fee and Long’s experimental methodology of cooling and heating bird brains, thereby effecting song production, and allowing inferences to be made on the neurological mechanisms. Such elaborate experimenting does nothing but pull us closer to understanding of bird songs, hopefully transferable in part to human language.

Prather and Mooney’s discussion links to the last chapter involving neurons, allowing for thought in this area to continue unabated. It is excellent to see that mirror neurons are being investigated in other species, as it is a very current topic with implications for language (e.g. Ramachandran 2011).

Beckers’ chapter is an interesting one that provides the fundamentals of bird vocalisation. She points out that different bird species have different types of vocalisations, needing further research. The question arises whether general similarities can be found across different bird species.

The article by Fisher provides a clear, systematic review on what the FOXP2 gene is, careful not to come to premature conclusions. What I found interesting was the coverage of a similar gene in mice, yet another species represented in this volume.

Scharff and Thompson present an all-encompassing review on FoxP2 in birds, and give hope to the future of its continued study. A good feature of this article is the covering of the variation of FoxP2 at different stages, showing the levels of FoxP2 expression affects the behaviour of birds -- which gives a good functional insight.

Ramus’ discussion on genes and dyslexia highlights their importance in such disorders, without forgetting environmental factors. While it is still yet unclear to which gene(s) are the main causes of dyslexia, slowly we are coming closer.

Fitch presents his review of Darwin’s theory on his 200th anniversary. Indeed, if one is a proponent of the musical origin of language, then Darwin rightfully deserves credit as Fitch argues. The theory itself on the other hand, is just one among many competing theories of language evolution.

Okanoya presents a series of reviews which build up a convincing array of evidence supporting his scenario presented. I appreciated the picture of the birds, and their historical background.

Pepperberg discusses her thoughts on mirror neurons in relation to humans and birds, most of which is speculative, but nonetheless deserves further attention. Moreover, as part of the articles here on mirror neurons, this again brings important focus on the phenomena which I believe holds plenty of promise for understanding the origins of language.

Evaluating the book as a whole, it is simply a splendid piece of scholarship. Firstly, the editors have done a superb job in collating such excellent scholarship from a range of authors who are clearly leaders in their respective fields. Without exception, all papers are well written, concise and clear to the point. The amount of different topics is also outstanding, including the number of different species covered. The editors are also to be commended in how the articles relate to one another, often on similar topics, with each article feeding off one another. Moreover, the amount of references the book contains will certainly assist those that are looking to enter the field. It is for this reason I would recommend the book to advanced undergraduates and beyond, where this book provides an excellent starting point.


Austin, Peter K. 2001. Word order in a free word order language: The case of Jiwarli. In: Jane Simpson, David Nash, Mary Laughren, Peter Austin and Barry Alpher (eds.), Forty years on: Ken Hale and Australian languages. Pp. 205-323. Canberra: Pacific Linguistics.

Hauser, Marc, Chomsky, Noam and Fitch, W. Tecumseh. 2002. The faculty of language: What is it, who has it, and how did it evolve? Science 298:1569-1579.

Huang, Yan. 2000. Anaphora. New York: Oxford University Press.

Lieberman, Philip. 2002. On the nature and evolution of the neural bases of human language. American Journal of Physical Anthropology, 119:36-62.

Pepperberg, Irene M. 2012. Further evidence for addition and numerical competence by a grey parrot (Psittacus erithacus). Animal Cognition 15(4):711-717.

Ramachandran, Vilayanur. S. 2011. The tell-tale brain: Unlocking the mystery of human nature. London: William Heinemass.

Wexler, Ken. 2011. Grammatical Computation in the Optional Infinitive Stage. In: Jill de Villiers and Tom Roeper (eds.), Handbook of Generative Approaches to Language Acquisition. Pp. 53-118. Dordrecht: Springer.

Yang, Charles. 2011. Computational Models of Language Acquisition. In: Jill de Villiers and Tom Roeper (eds.), Handbook of Generative Approaches to Language Acquisition. Pp.119-154. Dordrecht: Springer.


Dr. Darcy Sperlich is currently a senior lecturer of ESOL in the School of English at the Manukau Institute of Technology, in Auckland, New Zealand. His research interests include the study of pragmatic and syntactic theories in relation to anaphora (especially Chinese), experimental linguistic methodology, second language acquisition, and comparative Chinese syntactic dialectology.

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