Dieter G. Hillert- 1School of Speech, Language, and Hearing Sciences, San Diego State University, San Diego, CA, USA
- 2School of Medicine, University of California, San Diego, La Jolla, CA, USA
Introduction
Some language scientists defend an anti-Darwin account and believe in the saltational evolution of modern language. They emphasize that the language faculty emerged by a sudden mutation in the last 50–100 ky (e.g., Klein, 2000; Chomsky, 2012, 2015; Berwick et al., 2013). In contrast, others claim that modern language is the product of a gradual co-evolution of neurobiological and cultural-linguistic conditions, which took place since genus Pan was separated for good from the hominin lineage about 4–6 mya (e.g., Pinker and Bloom, 1990; Pinker, 1994; Deacon, 1997; Dor and Jablonka, 2001; Falk, 2004; Enfield and Levinson, 2006; Levinson and Jaisson, 2006; Christiansen and Chater, 2008; Atkinson, 2011; Dunn et al., 2011; Dediu and Levinson, 2014). New genetic evidence and their interpretation in context of fossil and artifact discoveries shed however light on this controversy. The data indicate that pre-modern language might have been already spoken by Homo erectus. Moreover, we conclude that the sister species of modern humans, Neanderthals and Denisovans, may have used language much like modern humans do (e.g., Dediu and Levinson, 2013).
To begin with, it is important to distinguish between the biology of modern language, that is, the language-ready brain and the availability of a linguistic code. For instance, a particular Homo species might have had a (pre-)modern language-ready brain but the language that was used at that time may have been different from modern language. Basic design features including combinatory, compositional as well as complex hierarchical structures are cultural products, which may have co-evolved along with genetic changes over a long period of time.
Modern language as we experience it today may require not only cortical wiring for language-specific operations but also a certain cortical mass to express thoughts and complex concepts. It would go beyond the present scope to discuss details but we assume here that the complex properties of modern language at all linguistic levels are from an evolutionary viewpoint mutually related to the principles of complex concept formations. An increase of the cranial capacity resulted in forming multi-modal memory systems. It seems thus plausible to assume that spoken language processing skills evolved alongside in order to communicate these sensory and episodic experiences.
The dramatic brain growth started with Homo habilis, which is sometimes considered to have an anagenetic relationship to the species H. erectus (Spoor et al., 2007). Again, sometimes Homo heidelbergensis has been classified as part of the H. erectus group by taking account of a broad range of individual differences (Lordkipanidze et al., 2013). The species H. habilis was around for 1 my, H. erectus for 1.6 my and H. heidelbergensis for 400 ky. As we will see below, the H. erectus epoch might have provided the opportunity for gradual biological adaptions resulting in refined language use. Recent genetic studies revealed factors, which may have significantly contributed to brain growth in human ancestors. Interestingly, these genetic factors seem to correspond to discrete cognitive stages in the human lineage.
Genetic Mutations for Brain Growth
The human brain, whose size tripled compared to genus Pan in a period of approximately 4 my, is not the result of a single mutation but involves multiple mutations. These genetic changes were dependent on each other and these changes were mutually dependent and favored by natural selection. Multiple genes are involved in the growth of the human brain (Fortna et al., 2004), but recent studies show that two different genes, ARHGAP11B and SRGAP2, seemed to have played a major role in cortical (re)organization in hominins. Brain growth for the evolution of modern language at all linguistic levels is crucial.
The gene ARHGAP11B was partially copied from ARHGAP11A after the hominin lineage split for good from genus Pan. As far as known, this copy exists only in modern humans, Neanderthals and Denisovans. In using an embryonal mouse model, Florio et al. (2015) showed that ARHGAP11B has the highest degree of radial glia-specific expressions. It promotes basal progenitor generation and proliferation, increases the cortical plate area and induces gyrification. ARHGAP11B can induce the folding of the developing mouse neocortex and may have contributed to the expansion of the human neocortex and their biological ancestors.
Also, the gene duplicates of SRGAP2 significantly contributed to the expansion of the neocortex while the transition from Australopithecus to Homo took place. Dennis et al. (2012) found a copy-code mechanism, that is, in addition to the ancestral duplications at chromosome 1 further copies of SRGAP2 were discovered. Thereby, the copies are not complete duplicates but are missing a small piece of the ancestral gene. The first duplications SRGAP2A and SRGAP2B were created ca. 3.2 mya and SRGAP2C was copied from SRGAP2B about 2.4 mya. Finally, the copy SRGAP2D appeared 900 kya, also a mutation of SRGAP2B. In particular SRGAP2C seemed to have accelerated cortical connectivity as indicated by mouse models. Charrier et al. (2012) found in knockdown mice that the ancestral SRGAP2 will be mimicked by SRGAP2C and that neurons travel faster to the target areas as compared to neurons without the C variant. The inhibition of SRGAP2 by human-specific paralogs certainly has played an important role in human brain development and thus presumably for the development of complex cognitive abilities. The occurrence of particular SRGAP2 copies seems to correspond to some extent to certain epochs, in which new hominins with larger brains evolved and repopulated our world.
Australopithecus lived in Africa between 4 and 2 mya and had an average brain size of 450 cc, which is comparable to the size of chimpanzees. In this epoch, ca. 3.4 mya, the mutations SRGAP2A and SRGAP2B occurred. In particular SRGAP2C, a copy of SRGAP2B, significantly contributed, about 2.4 mya, to the growth of the human neocortex. The dating of the fossils classified as H. habilis (2.4–1.4 mya) and H. erectus (1.9–0.3 mya) corresponds to the occurrence of this genetic copy mechanism. On average, H. habilis had had a cranial capacity of 600 cc and H. erectus between 800 and 1000 cc. Finally, SRGAP2D appeared about 900 kya, which is as SRGAP2C a copy of SRGAP2B. The D-variant corresponds to the appearance of H. heidelbergensis or late H. erectus (200–600 kya). Typically, H. erectus is considered to be the direct ancestor of Neanderthals, Denisovans and modern humans. All three species co-existed and intermingled in Eurasia (e.g., Green et al., 2010; Meyer et al., 2012). Neanderthals lived throughout Europe and Middle East 28–200 kya, Denisovans fossils found in Siberia were dated to be 41 ky old and our species appeared in Africa about 200 kya and began to migrate to Eurasia ca. 60 kya. The brain volume of the Neanderthals was with 1 cc on average 250 cc larger than of modern humans. The Denisovans' brain size probably matched this range but because only one finger bone and two teeth were found (Krause et al., 2010), it is difficult to draw conclusions about their cranial capacity. In contrast to genus Pan, Neanderthals, Denisovans and modern humans obtained ARHGAP11B and the relevant SRGAP2 copies.
Although, the factor brain volume alone does not inform about neural connectivity and the type of cortical circuits, we can certainly state that during a period of 3 my SRGAP2 mutations played a major role in brain development. It is estimated that the adult brain of modern humans has ca. Twenty billion neocortical neurons and a single neuron has on average 7000 synaptic connections (Drachman, 2005; Herculano-Houzel, 2009). If the copy-code mechanism of SRGAP2 was significant for the development of cognitive abilities and thus presumably also for language, the hypothesis of a sudden mutation, which brought about the biology disposition of language, seems not to be a convincing hypothesis. Instead, a non-expressed latent mutation in genus Australopithecus has been the precondition for further expressed mutations to promote brain volume and reorganization in the lineage of genus Homo.
The biological disposition for cognition and language seems thus to be the result of a gradual mutation, which may have begun already 3 mya. This hypothesis is certainly not a conclusive presumption. We do not know at which point in time the biological disposition of (pre-)modern language evolved, but in considering the reported genetic data on the neocortical evolution, it seems that basic principles supporting modern language were biologically disposed much earlier than previously assumed (Dediu and Levinson, 2013).
From Vocalization to Spoken Language
Along with the evolving biological disposition of language linguistic knowledge must have developed. Since genetic data, fossils and artifacts do not inform us directly about social and linguistic behavior, the following assumptions are certainly speculative. Recently, however, the timeline for stone artifacts has been pushed back by about 700 ky (Harmand et al., 2015). The stone tools from Lomekwi 3 (West Turkana, Kenya), which are considered to be part of the Oldowan culture, are dated to be 3 my old. The only hominin species known to have lived in this region around this time is a Kenyan variant of Australopithecus afarensis. This species had an approximate brain size of 430 cc, which is comparable to the brain size of genus Pan. Apparently, Australopithecus made already use of an elaborated hand-motor control, which might indicate a reorganization of cortical structures as compared to genus Pan (Stout et al., 2008; Stout, 2010; Stout and Chaminade, 2012). Accordingly, perhaps Australopithecus was already able to use non-symbolic, but referential vocalizations (possibly in combination with facial expressions and gestures) to display basic emotions and perceptions.
During the H. erectus epoch, Oldowan tools were refined and gradually replaced by Acheulean tools, which indicates that along with the increase of the cranial capacity cognitive skills co-evolved. With the emergence of H. erectus (brain volume of ca. 1000 cc), the cranial capacity consisted not only of more neurons but also of more interconnections between neurons and different cortical areas to support cognitive processing. The invention of a linguistic code for words and phrasal structures might have been happened during the H. erectus epoch. This time window permitted a continuous cultural development.
Here, we do not discuss how basic vocalizations converged into phonological units with iconic and symbolic meanings or how lexical units were combined to phrases and sentences with a specific word order to express more complex meanings. The cultural-linguistic process certainly co-evolved gradually, but we do not know how many cultural steps were involved before modern language was created (e.g., Boyd and Richerson, 1985; Evans and Levinson, 2009). The descendants of H. erectus are the sister species Neanderthals and Denisovans, and modern humans. Presumably, the neocortices of Neanderthals and Denisovans were biologically similar to modern humans. For instance, Neanderthals seem to have undergone the same selective sweep for the speech related gene FOXP2 as modern humans in comparison to genus Pan (Enard et al., 2002; Krause et al., 2007; but see Coop et al., 2008) and similar results seem to apply for the genetic base of auditory capabilities (Martínez et al., 2004).
Conclusions
The viewpoint that the language faculty emerged as a result of a sudden mutation 50–100 kya, seems not to be well supported. As partly discussed, the black box of the evolution of cognitive systems is not completely opaque anymore. Genetic data, fossils, and artifacts clearly inform us about a possible evolutionary path, which resulted in cortical rewiring and modern language processing as known to us today. From a methodological viewpoint it is essential to differentiate between the biological and cultural-linguistic evolution. Here, we believe that H. erectus and possibly its related (sub)species might have invented fundamental structures as found today in modern languages. In considering current genetic evidence about the cortical development in the hominin lineage, there is no reason not to believe that the known sister species Neanderthals and Denisovans (and at present unknown related species) used spoken language much like modern humans do.
Conflict of Interest Statement
The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
References
Atkinson, Q. D. (2011). Phonemic diversity supports a serial founder effect model of language expansion from Africa. Science 332, 346–349. doi: 10.1126/science.1199295
Berwick, R. C., Hauser, M. D., and Tattersall, I. (2013). Neanderthal language? Just-so stories take center stage. Front. Psychol. 24:671. doi: 10.3389/fpsyg.2013.00671
Boyd, R., and Richerson, P. J. (1985). Culture and the Evolutionary Process. Chicago, IL: The University of Chicago Press.
Charrier, C., Joshi, K., Coutinho-Budd, J., Kim, J.-E., Lambert, N., de Marchena, J., et al. (2012). Inhibition of SRGAP2 function by its human-specific paralogs induces neoteny during spine maturation. Cell 149, 923–935. doi: 10.1016/j.cell.2012.03.034
Chomsky, N. (2012). The Science of Language. Interviews with James McGilvray. Cambridge, MA: Cambridge University Press.
Chomsky, N. (2015). Some core contested concepts. J. Psycholinguist. Res. 44, 91–104. doi: 10.1007/s10936-014-9331-5
Christiansen, M. H., and Chater, N. (2008). Language as shaped by the brain. Behav. Brain Sci. 31, 489–509; discussion 509–58. doi: 10.1017/s0140525x08004998
Coop, G., Bullaughey, K., Luca, F., and Przeworski, M. (2008). The timing of selection at the human FOXP2 gene. Mol. Biol. Evol. 25, 1257–1259. doi: 10.1093/molbev/msn091
Deacon, T. W. (1997). The Symbolic Species: The Co-evolution of Language and the Brain. London: Penguin Books.
Dediu, D., and Levinson, S. C. (2013). On the antiquity of language: the reinterpretation of Neandertal linguistic capacities and its consequences. Front. Psychol. 4:397. doi: 10.3389/fpsyg.2013.00397
Dediu, D., and Levinson, S. C. (2014). “The time frame of the emergence of modern language and its implications,” in The Social Origins of Language, eds D. Dor, C. Knight, and J. Lewis (Oxford: Oxford University Press), 184–195.
Dennis, M. Y., Nuttle, X., Sudmant, P. H., Antonacci, F., Graves, T. A., Nefedov, M., et al. (2012). Evolution of human-specific neural SRGAP2 genes by incomplete segmental duplication. Cell 149, 912–922. doi: 10.1016/j.cell.2012.03.033
Dor, D., and Jablonka, E. (2001). From cultural selection to genetic selection: a framework for the evolution of language. Selection 1, 33–56. doi: 10.1556/select.1.2000.1-3.5
Drachman, D. A. (2005). Do we have brain to spare? Neurology 64, 2004–2005. doi: 10.1212/01.WNL.0000166914.38327.BB
Dunn, M., Greenhill, S. J., Levinson, S. C., and Gray, R. D. (2011). Evolved structure of language shows lineage-specific trends in word-order universals. Nature 473, 79–82. doi: 10.1038/nature09923
Enard, W., Przeworski, M., Fisher, S. E., Lai, C. S., Wiebe, V., Kitano, T., et al. (2002). Molecular evolution of FOXP2, a gene involved in speech and language. Nature 418, 869–872. doi: 10.1038/nature01025
Enfield, N. J., and Levinson, S. C. (2006). Roots of Human Sociality: Cognition, Culture and Interaction. Oxford: Berg Publishers.
Evans, N., and Levinson, S. C. (2009). The myth of language universals: language diversity and its importance for cognitive science. Behav. Brain Sci. 32, 429–448. doi: 10.1017/S0140525X0999094X
Falk, D. (2004). Prelinguistic evolution in early hominins: whence motherese? Behav. Brain Sci. 27, 491–503. doi: 10.1017/S0140525X04000111
Florio, M., Albert, M., Taverna, E., Namba, T., Brandl, H., Lewitus, E., et al. (2015). Human-specific gene ARHGAP11B promotes basal progenitor amplification and neocortex expansion. Science 347, 1465–1470. doi: 10.1126/science.aaa1975
Fortna, A., Kim, Y., MacLaren, E., Marshall, K., Hahn, G., Meltesen, L., et al. (2004). Lineage-specific gene duplication and loss in human and great ape evolution. PLoS Biol. 2:E207. doi: 10.1371/journal.pbio.0020207
Green, R. E., Krause, J., Briggs, A. W., Maricic, T., Stenzel, U., Kircher, M., et al. (2010). A draft sequence of the Neandertal genome. Science 328, 710–722. doi: 10.1126/science.1188021
Harmand, S., Lewis, J. E., Feibel, C. S., Lepre, C. J., Prat, S., Lenoble, A., et al. (2015). 3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya. Nature 521, 310–315. doi: 10.1038/nature14464
Herculano-Houzel, S. (2009). The human brain in numbers: a linearly scaled-up primate brain. Front. Hum. Neurosci. 3:31. doi: 10.3389/neuro.09.031.2009
Klein, R. G. (2000). Archaeology and the evolution of human behavior. Evol. Anthropol. 9, 17–36. doi: 10.1002/(SICI)1520-6505(2000)9:13.0.CO;2-A
Krause, J., Fu, Q., Good, J. M., Viola, B., Shunkov, M. V., Derevianko, A. P., et al. (2010). The complete mitochondrial DNA genome of an unknown hominin from southern Siberia. Nature 464, 894–887, doi: 10.1038/nature08976
Krause, J., Lalueza-Fox, C., Orlando, L., Enard, W., Green, R. E., Burbano, H. A., et al. (2007). The derived FOXP2 variant of modern humans was shared with Neanderthals. Curr. Biol. 17, 1908–1912. doi: 10.1016/j.cub.2007.10.008
Levinson, S. C., and Jaisson, P. (eds. 2006). “Evolution culture,” in A Fyssen Foundation Symposium. (Cambridge, MA: MIT Press).
Lordkipanidze, D., Ponce de Leòn, M. S., Margvelashvili, A., Rak, Y., Rightmire, G. P., Vekua, A., et al. (2013). A complete skull from Dmanisi, Georgia, and the evolutionary biology of early homo. Science 342, 326–331. doi: 10.1126/science.1238484
Martínez, I., Rosa, M., Arsuaga, J.-L., Jarabo, P., Quam, R., Lorenzo, C., et al. (2004). Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain. Proc. Natl. Acad. Sci. U.S.A. 101, 9976–9981. doi: 10.1073/pnas.0403595101
Meyer, M., Kircher, M., Gansauge, M.-T., Li, H., Racimo, F., Mallick, S., et al. (2012). A high-coverage genome sequence from an archaic Denisovan individual. Science 338, 222–226. doi: 10.1126/science.1224344
Pinker, S., and Bloom, P. (1990). Natural language and natural selection. Behav. Brain Sci. 13, 707–784. doi: 10.1017/S0140525X00081061
Spoor, F., Leakey, M. G., Gathogo, P. N., Brown, F. H., Antón, S. C., McDougall, I., et al. (2007). Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya. Nature 448, 688–691. doi: 10.1038/nature05986
Stout, D. (2010). The evolution of cognitive control. Topics Cogn. Sci. 2, 614–630. doi: 10.1111/j.1756-8765.2009.01078.x
Stout, D., and Chaminade, T. (2012). Stone tools, language and the brain in human evolution. Philos. Trans. R. Soc. Lond. B 367, 75–87. doi: 10.1098/rstb.2011.0099
Keywords: Homo erectus, ARHGAP11B, SRGAP2, cultural evolution, biological disposition, brain development, language evolution, Neanderthals
Citation: Hillert DG (2015) On the Evolving Biology of Language. Front. Psychol. 6:1796. doi: 10.3389/fpsyg.2015.01796
Received: 01 September 2015; Accepted: 06 November 2015;
Published: 24 November 2015.
Edited by:
Robin Clark, University of Pennsylvania, USAReviewed by:
Monica Tamariz, The University of Edinburgh, UKCopyright © 2015 Hillert. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Dieter G. Hillert, dhillert@ucsd.edu