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Frontiers in Evolutionary Neuroscience ·

Volume 4 - 2012 | https://doi.org/10.3389/fnevo.2012.00012

Birds, primates, and spoken language origins: behavioral phenotypes and neurobiological substrates

  • Christopher I. Petkov1,2*corresponding author
  • Erich D. Jarvis3*corresponding author
  1. 11 Institute of Neuroscience, Newcastle University Newcastle upon Tyne, UK
  2. 22 Centre for Behavior and Evolution, Newcastle University Newcastle upon Tyne, UK
  3. 33 Department of Neurobiology, Howard Hughes Medical Institute, Duke University Durham, NC, USA
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Abstract

Vocal learners such as humans and songbirds can learn to produce elaborate patterns of structurally organized vocalizations, whereas many other vertebrates such as non-human primates and most other bird groups either cannot or do so to a very limited degree. To explain the similarities among humans and vocal-learning birds and the differences with other species, various theories have been proposed. One set of theories are motor theories, which underscore the role of the motor system as an evolutionary substrate for vocal production learning. For instance, the motor theory of speech and song perception proposes enhanced auditory perceptual learning of speech in humans and song in birds, which suggests a considerable level of neurobiological specialization. Another, a motor theory of vocal learning origin, proposes that the brain pathways that control the learning and production of song and speech were derived from adjacent motor brain pathways. Another set of theories are cognitive theories, which address the interface between cognition and the auditory-vocal domains to support language learning in humans. Here we critically review the behavioral and neurobiological evidence for parallels and differences between the so-called vocal learners and vocal non-learners in the context of motor and cognitive theories. In doing so, we note that behaviorally vocal-production learning abilities are more distributed than categorical, as are the auditory-learning abilities of animals. We propose testable hypotheses on the extent of the specializations and cross-species correspondences suggested by motor and cognitive theories. We believe that determining how spoken language evolved is likely to become clearer with concerted efforts in testing comparative data from many non-human animal species.

Keywords

1. Introduction

Charles Darwin's theory on descent with modification as it applies to man (Darwin, 1871) had for many years been used to underscore the importance of non-human primates for unraveling the origins and neuronal precursors of spoken language (e.g., Hewes, 1973). Yet, in part because of the apparent lack of vocal learning or syntactic-like abilities in non-human primates, different camps have focused on either the differences between human and non-human primates or their similarities. This has resulted in many contentious debates on language evolution with regards to non-human primates (for reviews: Pinker, 1994; Hauser et al., 2002; Pinker and Jackendoff, 2005). Adding further complexity for understanding spoken language origins, recently the research focus has shifted towards species more distantly related to humans, such as certain groups of songbirds. This is in part because songbirds like humans and a few other species exhibit vocal learning and have what has been broadly classified as “syntactic-like” song production (Doupe and Kuhl, 1999; Jarvis, 2004; Bolhuis et al., 2010). A summary of a consortium on the origins of human language syntax and its biological foundations encapsulates some of the current thinking: Another area of agreement might seem surprising in light of many current “primate-centric” studies of language evolution (Burling, 2006; Hurford, 2007). Most participants felt that there were no true precursors of syntax to be found among our nearest relatives. For anything like a syntactic precursor one had to go as far afield as songbirds ….

Likewise, in a thought provoking essay, Bolhuis and Wynne (2009) questioned to what extent evolutionary theory can help us to understand cognitive brain mechanisms in living animals. Their perspective was illustrated by a cartoon depicting a scientist with the great hope of teaching a monkey to say “apple,” but realizing that the monkey is the classroom dunce when the parrot vocally identifies the apple variety as “golden delicious.” Darwin, however, would have likely filled the classroom with as many different animals as possible. In any case, the authors' conclusions are appropriately nuanced and seem to favor a broader comparative approach: “there is no a priori reason to assume that convergence will be more important than common descent or vice versa” (Bolhuis and Wynne, 2009).

We, as researchers that have studied non-human primates and birds, argue that the path toward understanding the origins of spoken language cannot be based on focusing on a few select species with or without communication abilities that are either thought to be most comparable to humans, or to reflect physiology most comparable to humans. Any “one animal centric” approach will only limit our capacity to unravel the evolutionary bases of spoken language. If for no other reason, without “other” species as points of reference, it would not be clear what is special about human communication. Moreover, a focus on certain species restrains the development of different animal model systems with distinct advantages for understanding the neurobiological mechanisms of human language-related processes, which is important for advancing treatment options for communication and language disorders. Thus, to better understand the origins of human spoken language we rely on a broad comparative approach that takes advantage of information obtained across animal taxa, letting each animal have their “say” on the question of language evolution. We are aware that to do so can only be achieved by additional comparative work that will require energy and investment, combined with efforts to stay objective, as best as we can, regarding the cross-species similarities and differences.

In an effort to invigorate a broader perspective on spoken language origins, in this paper we overview the parallels and differences in the behavioral and neurobiological data of vocal learners (e.g., humans and songbirds) and those animals often identified as “vocal non-learners.” We ask how strong is the evidence for categorical distinctions between vocal learners and vocal non-learners? We note that vocal non-learners are often classified as such based on a lack of experimental evidence, but that when the animals are tested, there is often more variation in vocal learning abilities than might have been expected (Janik and Slater, 2000; Snowdon, 2009; Arriaga et al., in press). Moreover, since vocal learning depends on auditory learning, and auditory learning abilities are broadly conserved in the animal kingdom, we ask how this trait dependency could have influenced the evolution and mechanisms of vocal learning. Then, based on a modified perspective of the literature we reconsider some of the motor and other theories that have been proposed for humans, birds and other animals. We conclude by generating testable hypothesis, including for: (1) better understanding variability in the vocal behavior and neurobiology of vertebrates that are often classified as vocal non-learners; and (2) the possible capabilities of, for example, non-human primates as limited vocal learners but considerable auditory learners, to learn the structure of auditory sequences, and whether this might tap into an ancestral “proto-syntactic” brain network that evolved in humans to support syntactic learning.

2. Conclusions

This review has considered the behavioral and neurobiological data in complex-vocal learners such as, humans and songbirds and how they relate to data from so-called “vocal non-learners.” We noted that the evidence provided by several recent examples in the animal behavioral literature motivates a revision of the hypothesized “vocal learning” vs. “vocal non-learning” distinction. We outlined an alternative hypothesis of greater variability in vocal learning categories and in a related but different behavioral phenotype, namely, auditory (sensory) sequence learning. Upon this modified perspective of the behavioral literature, we considered neurobiological distinctions between “vocal learners” and “non-learners,” questioning whether these distinctions will be useful for clarifying behavioral results or whether behavioral variability can help us to understand the neurobiological substrates and distinctions at a finer level.

Motor, gestural and other theories were considered and predictions made, including from the perspective of animals that are not complex-vocal learners. We also considered the distinctions between the tenets of a number of theories regarding the sensory learning (reception) and motor (production) systems, including interconnectivity and interactions with the neurobiological systems supporting cognitive processes and how these may have evolved. This was used to make specific predictions, exemplified in humans, birds, and non-human primates to investigate the neurobiological substrates that might be evolutionarily related, either by common descent or convergence, to the ones that humans rely on for language.

Taking all of this into consideration, we highlighted the need for comparative approaches that more closely consider the behavioral and neurobiological data on sensory learning and vocal production abilities in many vertebrates. This will be critical for staying objective, empirically grounded, and realistic with regards to the aspects of human spoken language that different animal species could address or serve to model. We hope that this paper has been able to underscore the importance of and to encourage further interdisciplinary cross-species work for clarifying not only the origins of spoken language but also the conserved components and specializations present in the neurobiological systems supporting the communication abilities of animals.

Conflict of interest statement

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Citation

Petkov, C. I. & Jarvis, E. D. (2012). Birds, primates, and spoken language origins: behavioral phenotypes and neurobiological substrates. Frontiers in Evolutionary Neuroscience, Volume 4. https://doi.org/
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*Correspondence: Christopher I. Petkov; Erich D. Jarvis

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