Annual Review of Genetics, 39, 197–218. Fossils, along with the comparative anatomy of present-day organisms, constitute the morphological, or anatomical, record. 143–156). The evolution of vocal learning. Berlin, Germany: Springer-Verlag. This set of components—the so-called faculty of language in a broad sense—is large, but divides naturally into two categories: those that are shared (sometimes widely) with other species and those that are recent acquisitions of the human lineage since our evolutionary divergence from chimpanzees. In E. H. Lenneberg & E. Lenneberg (Eds. ), Language, music, and the brain: A mysterious relationship (Vol. Denton, M. (1985). Evolutionary linguists consider linguistics as a subfield of evolutionary biology and evolutionary psychology. The now classic case concerns complex eyes in insects and vertebrates, which were not present in the common ancestor (and are phenotypically analogies) but nonetheless rely upon many of the same regulatory genes (e.g., Pax-6) for their development (genetic homologies). The earth is round (p <.05). Especially when an attempt is made to have the hypothesis set tested be exhaustive, this method has been dubbed “strong inference” (Chamberlin, 1890), and when thoughtfully implemented offers a much more efficient path to resolution of scientific debates and apparently discrepant data. Although the correct theoretical and philosophical framework for understanding this picture remains a topic of discussion (see, e.g., the articles by Arbib, 2016; Chomsky, 2016; Jackendoff & Wittenberg, 2016; Scott-Phillips, 2016), it all concerns modern humans and thus defines the “end target” of any evolutionary model, rather than the steps required to get to this point. Nonetheless, the belief remains distressingly widespread that major changes in vocal anatomy, and particularly the descended human larynx, were required prerequisites for speech. Similar to musical protolanguage, Donald envisions a crucial role for performative, group-defining rituals, initially devoid of propositional meaning, as an initial stage of language evolution, one he ties to Homo erectus. (2003). Proceedings of the Royal Society of London B, 268(1477), 1669–1675. Social cognition and the evolution of language: Constructing cognitive phylogenies. Thus, the key innovation at this stage is dendrophilia (Fitch, 2014)—a domain-general proclivity to perceive hierarchical structure that came to be applied not only to language but also to music and decorative arts. Once an elaborate, socially shared but nonpropositional communication system is in place, the major problem is how propositional semantics could be added to such a system. Nature Communications, 5, 483. But unfortunately, finding further clear differences will involve both luck and hard work, because there has been so much genetic change that it is difficult to come to grips with. Language learning: Language is transformed as it is transmitted from one generation to the next. The antidote to this persistent problem is to acknowledge Deep homology and the origins of evolutionary novelty. Phillip Lieberman has investigated the origin of speech for many years and has used this research to form hypotheses about the evolution of language. Mesoudi, A., Whiten, A., & Laland, K. N. (2004). This is what Chomsky calls “the creative aspect of language use”. Punctuated equilibrium—A different way of seeing. 119–130). (2005). (2012). Nature, 212, 1468–1469. While such complex repertoires are far from trivial, and their strings differ discriminably from one another, they do not communicate equally complex semantic messages. Berlin, Germany: Druckerei der Königlichen Akademie der Wissenschaften. Typically the birth of modern hierarchical syntax is seen as the final stage of language evolution, so this is a “syntax last” model. This process has led, for example, to plate tectonics going from a speculative hypothesis, often ridiculed, to something universally accepted in modern geology (Gohau, 1990). Kroodsma, D. E. (2005). Evolution of vertebrate colour vision. Hublin, J.-J. The human tongue is similar in anatomy to that in other apes (Takemoto, 2008), and we now know that a mild descent of the larynx and hyoid bone occurs in chimpanzees (Nishimura, Mikami, Suzuki, & Matsuzawa, 2006). Hancock, A. M., & Di Rienzo, A. The final general point concerns the need for simultaneous testing of multiple, plausible hypotheses. Language & Communication, 18(1), 47–67. Theissen, G. (2009). Pinker, S., & Bloom, P. (1990). Representation of numerical and sequential patterns in macaque and human brains. (2013). In N. L. Wallin, B. Merker, & S. Brown (Eds. Cambridge, MA: MIT Press. This combination has led to increasingly sophisticated models of language evolution that make multiple testable predictions, and improved evaluation criteria for assessing such models. Interest in language evolution has surged in the past two decades (Berwick, Friederici, Chomsky, & Bolhuis, 2013; Carstairs-McCarthy, 1999; Christiansen & Kirby, 2003; Dunbar, 1996; Fitch, 2010; Hauser, Chomsky, & Fitch, 2002; Pinker & Bloom, 1990; Pinker & Jackendoff, 2005; Számadó & Szathmary, 2006; Tallerman & Gibson, 2011; Yang, 2013). Working memory of numerals in chimpanzees. Shaping of hooks in New Caledonian crows. Premack, D., & Woodruff, G. (1978). Endicott, P., Ho, S. Y. W., & Stringer, C. (2010). This surge has been accompanied by fundamental progress in our understanding of this difficult multidisciplinary problem. Behavioral and Brain Sciences, 4, 515–526. The central quantitative measure of selection in comparisons between species is the dN/dS ratio: the ratio of nonsynonymous to synonymous base pair changes. doi:10.3758/s13423-016-1101-y. Bugnyar et al., 2016; Call & Tomasello, 2008; Fitch et al., 2010; Scott-Phillips, 2016; Seyfarth & Cheney, 2016), where the cognitive parallels of pragmatics and theory of mind can clearly be found in ravens. Patel, A. D. (2016). Lankester, E. (1870). 226–271). Also, if we accept that some capacities key to language may have evolved in contexts other than communication (and were later exapted during language evolution), then we need to consider data from animal cognition on an equal footing with animal communication. Language, 87(3), 586–624. Clearly, language as a system for expressing thought is not limited to the audio-vocal channel (for further implications of this fact, see de Boer, 2016; Goldin-Meadow, 2016; Kendon, 2016). A host of other auditory phenomena such as categorical perception or vowel “prototype magnets” have also been documented in other species from birds to chinchillas (Kluender, Diehl, & Killeen, 1987; Kuhl & Miller, 1978). Another trait that appears to differentiate humans from other apes concerns not our ability to communicate, but rather our proclivity to do so, a proclivity for which I have borrowed the German word Mitteilungsbedürfnis (the drive to share thoughts). The ecology and evolution of primate color vision. Boyd, R., & Richerson, P. J. Phonetica, 57, 205–218. ), Foundations of language development: A multidisciplinary approach (Vol. Donald’s piece in the current issue lays out this model and predictions concisely. MacLarnon, A. M., & Hewitt, G. P. (1999). In D. Scarborough & S. Sternberg (Eds. Thus, there is little reason today to accept the old assertion that “speech is special” from a perceptual point of view (Liberman, Cooper, Shankweiler, & Studdert-Kennedy, 1967), or to think that new mechanisms of speech perception needed to evolve in the hominid lineage to support language evolution. Auk, 94, 783–785. This notion of change of function is a central, although often overlooked, plank of evolutionary thought (Pievani & Serrelli, 2011) because it can refute arguments of “irreducible complexity” (Denton, 1985)—that complex organs like wings or eyes could not have evolved from simple beginnings because they need to be complex to have any functionality at all. The first is that the primary determinant of hearing range and acuity is the cochlea, not the outer or middle ears (Ruggero & Temchin, 2002). (Eds.). Scott-Phillips, T. C. (2016). Wild, J. M. (1997). Psychonomic Bulletin & Review. The signature of positive selection at randomly chosen loci. The mystery of language evolution. In fact, there were several scientists that came before Darwin that had hypothesized the exact same thing. Hurford, J. The Cambridge encyclopedia of language (2nd ed.). London, UK: Weidenfeld & Nicolson. This is one major way researchers reconstruct the evolution of language — by studying if and to what degree (nonhuman) animals possess its requisite parts. It is certainly not the capacity to produce rule-governed behavior, nor a capacity to produce or interpret signals—virtually any vertebrate does these things. Tongue protrusions provide a new method to study language evolution and development. Thus in the same way that today's neuroscientists focus less on individual neurons and more on neuronal circuits (and less on specific brain regions than on global brain functional networks), the new genomic era forces us to think more in terms of genetic circuits, gene regulation, and interactions among multiple alleles than was typical during the pregenomic era. (1974). Hillsdale, NJ: Erlbaum. Trends in Cognitive Sciences, 7(5), 187–189. Physical production mechanism of elephant infrasonic vocalizations. PubMed ), Darwin up to date (Vol. But I hope it clarifies the essential point that most of the mechanisms involved in language acquisition evolved long before humans split with chimpanzees. Neuron, 88, 2–19. Udden, J., Folia, V., Forkstam, C., Ingvar, M., Fernandez, G., Overeem, S., van Elswijk, G., … Petersson, K. M. (2008). European Journal of Human Genetics, 22, 171–178. Not necessarily a wing: Which came first, the function or the form? Learn vocabulary, terms, and more with flashcards, games, and other study tools. Loss of function of a particular protein led to inactivation of a muscle protein expressed particularly in the temporalis jaw muscle, MYH16, and may have led to the reduction in this muscle’s size and jaw robusticity (Stedman et al., 2004). Boston, MA: Houghton Mifflin. Musical protolanguage models also suggest a close link between speech phonology and song, a prediction that can be tested using both brain imaging and genetic data. Systematic Zoology, 22, 425–441. Stoeger, A. S., Mietchen, D., Oh, S., de Silva, S., Herbst, C. T., Kwon, S., & Fitch, W. T. (2012). The current list of gifted vocal learners includes some species of songbirds, hummingbirds, parrots, cetaceans, seals, bats, and elephants. American Journal of Radiology, 178, 3–16. Perfors, A. As a member, you'll also get unlimited access to over 83,000 lessons in math, English, science, history, and more. National Geographic Headquarters While there have also been naysayers who deny this progress, or the very possibility of progress in understanding cognitive evolution (Hauser et al., 2014; Lewontin, 1998), the purpose of this review and the special issue of which it is part is to demonstrate that such pessimism is unjustified. Artificial grammar learning meets formal language theory: An overview. Philosophical Transactions of the Royal Society, B: Biological Sciences, 361, 23–43. I will refer to these core traits derived components of language (relative to our last common ancestor with chimpanzees), or “DCLs.” By our current understanding of ape cognition and communication, the set of DCLs contains at least three separable components (see The derived components section): complex vocal learning, hierarchical syntax, and complex semantics/pragmatics (cf. The answer initially is “no one” (at least until offspring were born who shared the mutation), but the ability would nonetheless be useful in private thought. Evolution of the neural language network. Phylogeny, the history of the evolution of a species or group, especially in reference to lines of descent and relationships among broad groups of organisms. I want to underscore the importance of using the term "evidence" rather than the more colloquial term 'proof' in normal scientific discourse. Start studying 2.07 Quiz: Evidence for Evolution 2. Localisation of a gene implicated in a severe speech and language disorder. Cosmologists interested in the Big Bang, or geologists studying continental drift and plate tectonics, are quite familiar with such difficulties. B) Evolution of the brain occurred first leading to language development. Cambridge, MA: MIT Press. Herrmann, Call, Hernàndez-Lloreda, Hare, & Tomasello, 2007). It is easy to overlook the importance of this trait, but without it the free flow of information that provides a prime benefit of language would slow to a trickle. An RNA gene expressed during cortical development evolved rapidly in humans. Catani, M., & Mesulam, M. (2008). Philosophical Transactions of the Royal Society B, 366(1574), 2124–2140. The symbolic species: The co-evolution of language and the brain. (In this issue). We also have high-quality genomes for two extinct archaic hominins, Neanderthals and Denisovans. Decreased activity of this inhibitory gene may be one of several changes involved in the expansion of brain size during hominin evolution. Przeworski, M. (2002). Proceedings of the National Academy of Sciences. Fitch, W. T. (2013). (2012). New York, NY: Academic Press. Over thousands of years, humans have developed a wide variety of systems to assign specific meaning to sounds, forming words and systems of grammar to create languages. As these examples make clear, our understanding of cognitive evolution would be seriously incomplete if we focused exclusively on comparisons of humans with other primates (a narrow comparative approach). Gould, S. J. Join the dots: A musical interlude in the evolution of language? Language-tree divergence times support the Anatolian theory of Indo-European origin. Monographs of the Society for Research in Child Development, 58, 1–221. We have a relatively clear endpoint of the process in the present, and can reconstruct the starting point (our last common ancestor with chimpanzees) in detail using the comparative method with existing species. Nottebohm, F. (1972). Left hemisphere specialization for oro- facial movements of learned vocal signals by captive chimpanzees. What young chimpanzees know about seeing. How can we detect when language emerged? The roles of the external, middle, and inner ears in determining the bandwidth of hearing. But it is now clear that even RNA that is not translated into protein can, by itself, play myriad biological roles, and this HARF1 gene is an interesting example, again involved in brain development. New York, NY: Cambridge University Press. Turning to signal output, there are no clear differences between humans and other apes in terms either of vision or manual/facial control that would prevent an ape from learning signed language if the proper semantic and syntactic “software” were in place—one reason for the popularity of “gestural protolanguage” hypotheses (see below and Corballis, 2002; Hewes, 1973). to the clear recognition of the importance of brain networks (often spread widely throughout cortex) and connectivity in neural computation. The human infant brain: A neural architecture able to learn language. Philosophical Transactions of the Royal Society B, 367, 88–102. (1996). Since Darwin’s time, scientists have puzzled over the evolution of language.They can observe the present-day product: People today have the capacity for language, whether it be spoken, signed or written. (2016). The core idea of the Minimalist paradigm in linguistics is that linguistic syntax can be reduced to a very simple but powerful core operation, Merge, which serves to combine lexical elements (Chomsky, 1995); this conception opened the door to inquiry into the evolution of such an operator (Chomsky, 2010, 2016). All of these models imply that syntax was a very late acquisition in language evolution. Lyn, H. (2017). Vision: A computational investigation into the human representation and processing of visual information. Recent data from chimpanzees, where food calling behavior of newcomers to a zoo population gradually became more similar to that of previous residents (Watson et al., 2015), show nothing more than this type of call usage learning (and are not particularly convincing evidence of that; Fischer, Wheeler, & Higham, 2015). ), The origins of music (pp. Oxford, UK: Oxford University Press. Shea, J. J. Jacob, F. (1977). Concepts such as adaptation and mutation inform therapies and strategies to combat pathogens, including influenza. https://doi.org/10.3758/s13423-017-1236-5, DOI: https://doi.org/10.3758/s13423-017-1236-5, Over 10 million scientific documents at your fingertips, Not logged in Trends in Cognitive Sciences, 12(5), 187–192. More specialized capacities including tool use are also shared, not only with other primates but with a variety of avian and mammalian species (McGrew, 2004; Pruetz & Bertolani, 2007; Tebbich, Taborsky, Fessl, & Blomqvist, 2001; van Lawick-Goodall & van Lawick-Goodall, 1967; Weir, Chappell, & Kacelnik, 2004; Whiten, Horner, & de Waal, 2005). Psychonomic Bulletin & Review. doi:10.3389/fpsyg.2014.00401. Instead the tradition has been one where others’ models are ridiculed or (worse) ignored. Nature, 399, 682–685. My aim here is illustrative, to show that a model can be constructed that is consistent with all available data, and that makes clear testable predictions. Many organisms can also make “infinite use of finite means” in the restricted sense that they can produce an unlimited variety of signal strings (think of an incessantly barking dog—no day’s barking will be precisely the same as another’s). London, UK: IPC Magazines. Fitch, W. T., & Jarvis, E. D. (2013). Harvey, P. H., & Pagel, M. D. (1991). Language evolution. As Darwin pointed out, the forelimbs of such animals as humans, porpoises, bats, and other creatures are strikingly similar, even though the forelimbs are used for different purposes (that is, lifting, swimming, and flying, respectively). What was the trigger and chain of events that caused such a massive upset of Earth's biota? Of these four classes of data, the most exciting are genetic data, and particularly paleo-DNA from extinct hominins, which offer the tantalizing hope of explicitly testing and rejecting predictions of current models of language evolution. Journal of Neuroscience, 32(41), 14125–14131. Nature, 418, 869–872. The effects of deep-reaching lesions in the cortical face area on phonation: A combined case report and experimental monkey study. Primate precursors to human language: Beyond discontinuity. Watson, S. K., Townsend, S. W., Schel, A. M., Wilke, C., Wallace, E. K., Cheng, L., … Slocombe, K. E. (2015). Pascual-Leone, A., Walsh, V., & Rothwell, J. But for no one of these subcomponents do we have clear evidence of a definitive qualitative difference: it seems to be the whole package that evolved. These are two related species of humans, whose homelands were Europe and Asia, that evolved independently of modern Homo sapiens, in these regions. The evolution of languages has been happening for hundreds if not thousands of years, and it is natural. There is a growing consensus, well reflected in this issue, that cultural change has an important part to play in understanding language (Adger, 2017; Bowling, 2017; Kirby, 2017; Pagel, 2016; Steels, 2016;). (1973). This research has recently taken a strong empirical turn both in humans (Kirby et al., 2008; Morgan et al., 2015; Smith & Kirby, 2008) and animals (Fehér, 2016; Fehér, Wang, Saar, Mitra, & Tchernichovski, 2009; Whiten et al., 1999). Inhibition of SRGAP2 function by its human-specific paralogs induces neoteny during spine maturation. (2003). Oxford, UK: Oxford University Press. These results indicate that a capacity for first-order ToM was already present in the LCAc. Gyger, M., & Marler, P. (1988). Finally the fifth “Synthesis” part of the article attempts to do something I have resisted previously: It offers a comprehensive, testable model of language evolution, from our last common ancestor with chimpanzees to modern humans. Psychonomic Bulletin & Review. These new data also offer unprecedented ways to test hypotheses about language evolution and to uncover the timeline through which genes specifically involved in various DCLs changed and spread through our species. Baltimore, MD: Lippincott Williams & Wilkins. A chimpanzee recognizes synthetic speech with significantly reduced acoustic cues to phonetic content. Donald, M. (1991). By studying this type of evidence, archeologists can understand how early humans made and used tools and lived in their environments. But, as I will argue below, it will be worth it, because genes provide the closest thing to “fossils of language” we will ever have, and paleo-DNA in particular provides the analog of a time machine, taking us back nearly half a million years to the split between early Neanderthals and modern humans. Gould, S. J., & Eldredge, N. (1977). This introduction offers an overview of the field, and a summary of what needed to evolve to provide our species with language-ready brains. It is now clear that one of the first events in the hominin lineage was the habitual assumption of bipedal posture, and that this occurred before any major increase in brain size (Stringer & Andrews, 2005). Current Opinion in Neurobiology, 28, 48–53. Current Biology, 10(12), 723–726. This approach yields the strongest signal in cases where a particular region of the genome has been under continued selection, for example, in regions involve in disease resistance. confound the language of all the earth,” as described in Genesis (11: 9). Over the years, primate communication research has taken different forms. Another theme of the genomic revolution is the importance of gene duplication in evolution. Although models like these are often termed “theories” of language evolution, I prefer to use the term “model” because “theory” in science typically connotes a much more well-tested and widely accepted model than any existing models of language evolution. Washington, DC 20036, National Geographic Society is a 501 (c)(3) organization. The first concerns the internal (thought) versus external (communication) uses of language. The four sub-fields that comprise generative grammar include: phonology (the study of how languages sound), morphology (the study of how words are formed and what they mean), syntax (the study of the structure of sentences), and semantics (the study of linguistic meaning). doi:10.3758/s13423-016-1089-3. Overwhelming evidence supports this fact. But simplicity at the computational level of description does not necessarily translate into implementational simplicity at the neural level (or vice versa), and simplicity of neural implementation is arguably the level most relevant to evolutionary discussions of rapid adaptive change (Johnson, 2016; Perfors, 2017). Savage-Rumbaugh, E. S. (1986). The human capacity for complex vocal learning is a case in point: though basically absent in chimpanzees or other primates (see below), it is shared with a diverse if scattered collection of other bird and mammal species (Brainard & Fitch, 2014; Fitch & Jarvis, 2013; Janik & Slater, 1997). How protolanguage became language. Atypical birdsong and artificial languages provide insights into how communication systems are shaped by learning, use, and transmission. doi:10.1073/pnas.1605782113. Psychonomic Bulletin & Review. San Diego: Academic Press. The complete genome sequence of a Neanderthal from the Altai Mountains. Darwin, C. (1871). doi:10.3758/s13423-016-1061-2. Psychonomic Bulletin & Review. This special issue provides a very rich selection of comparative data, including work on nonhuman primates (Arbib, 2016; Byrne & Cochet, 2016; Fischer, 2016; Lyn, 2017; Seyfarth & Cheney, 2016), birds (Fehér, 2016; Okanoya, 2017; ten Cate, 2016), bats (Vernes, 2016), or a mixture of species (Pepperberg, 2016). While the dataset for comparative comparisons is sparser, we have a rather clear conception of what precisely changed during the evolution of the human brain: In addition to a general size expansion, there were particular expansions (both in raw size and in terms of connectivity) of brain regions long known to play an important role in linguistic syntax. They mark the third crucial domain of difference between humans and our nearest primate relatives (Herrmann et al., 2007). Deacon, T. W. (1992). During this “singing ape” stage, sequencing of learned signals was already in place, adding a selective pressure for rapid and accurate sequence learning as typified by many modern songbirds (Jarvis, 2004b; Kroodsma, 2005; Marler & Slabbekoorn, 2004). The existence of multiple DCLs leads logically to a notion of “protolanguage”—some hypothesized system of thought and/or communication that had some DCL(s) but not the full suite. Kuhl, P. K., & Miller, J. D. (1978). Selective scenarios for the emergence of natural language. (2013). A potentially language-relevant example of the first phenomenon is SRGAP2, a gene which has duplicated three times in humans relative to other apes. Fitch, W. T. (2011e). A programming language is an artificial language that can be used to control the behavior of computer. Psychonomic Bulletin & Review. American Zoologist, 40, 862–873. Finally, one of the classic supposed differences between humans and other apes was our possession of a “theory of mind”—a capacity to represent the beliefs and desires of other individuals (Povinelli & Eddy, 1996; Premack & Woodruff, 1978). Gathering evidence to study mass extinctions. Foundations of language. Elder, J. H. (1934). Holloway, R. L., Broadfield, D. C., Yuan, M. S., Schwartz, J. H., & Tattersall, I. Cambridge, MA: MIT Press. Kendon, A. Oxford, UK: Oxford University Press. Journal of Archaeological Science, 20, 81–91. below and Hurford, 1990). In J. Blasi, D. E., Wichmann, S., Hammerström, H., Stadler, P. F., & Christiansen, M. H. (2016). Poletiek, F. H., Fitz, H., & Bocanegra, B. R. (2016). In P. Marler & H. Slabbekoorn (Eds. Believing this account, however, requires believing in God, and the denial of the evolution theory, which suggests that all animals, humans, and even human language, arose by chance. Mithen, 2005). Recent research using artificial grammar learning to probe the specific types of rules that organisms are capable of learning has offered a clearer picture (see ten Cate, 2016), and the use of formal language theory provides a useful metric and common description language for analyzing these rule types in terms of computational complexity (Jäger & Rogers, 2012). Fitch, W. T., Hauser, M. D., & Chomsky, N. (2005). This area is greatly expanded in humans relative to chimpanzees (Schenker et al., 2010). The evolution of syntax: An exaptationist perspective.
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