modern evolutionary thinking in language evolution and … · modern evolutionary thinking in...

Dan Dediu 1

24th of July, 2014Euro Evo-Devo 2014Vienna, Austria

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Modern evolutionary thinking inlanguage evolution and change

Dan Dediu

Language and GeneticsMax Planck Institute for Psycholinguistics

Nijmegen, The Netherlands

Articulation of /r/Neandertal man Linguistic diversity

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The mystery of language evolution*

* Hauser, M. D., Yang, C., Berwick, R. C., Tattersall, I., Ryan, M., Watumull, J., … Lewontin, R. (2014). The mystery of language evolution. Frontiers in Psychology 5:401.

Present~ 2 mya~ 6 mya

MRCA

H. erectus

Neandert(h)als

Denisovans

Chimps

Language

No language

Very probablyno language

The “mystery”

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Broad models for language origins

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012 mya

H. erectus

Neandertals

Modern humans

Denisovans

~0.5 ~0.25

~0.03

H. heidelbergensis

Africa

Outside Africa

PNG, Australia...

Eurasia...

Reich, D., Patterson, N., Kircher, M., Delfin, F., … Stoneking, M. (2011). Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania. American Journal of Human Genetics 89:516–528.

Meyer, M., Kircher, M., … Pääbo, S. (2012). A High-Coverage Genome Sequence from an Archaic Denisovan Individual. Science 338:222–226.

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1

012 mya

H. erectus

Neandertals

Modern humans

Denisovans

~0.5

H. heidelbergensis

Klein, R. G. (2009). The Human Career: Human Biological and Cultural Origins (3rd edition). University of Chicago Press.Chomsky, N. (2010). Some simple evo devo theses: how true might they be for language. The evolution of human language: biolinguistic perspectives, 62:54–62.

Modern language

“Proto-language”

The “standard” view: language emerged recently and abruptly

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1

012 mya

H. erectus

Neandertals

Modern humans

Denisovans

~0.5

H. heidelbergensis

Dediu, D., & Levinson, S. C. (2013). On the antiquity of language. Frontiers in Language Sciences, 4:397.Frayer, D. W., Fiore, I., Lalueza-Fox, et al. (2012). Replay to Benítez-Burraco & Longa: When is enough, enough? Journal of Anthropological Sciences 90:1–6.D’ Errico, F., & Vanhaeren, M. (2009). Earliest personal ornaments and their significance for the origin of language debate. In R. Botha & C. Knight (Eds.) (pp. 16–40).

Modern language

“Proto-language”

The “alternative” view: language emerged early and piecemeal

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1Dediu, D., & Levinson, S. C. (2013). On the antiquity of language. Frontiers in Language Sciences, 4:397.Johansson, S. (2013). The Talking Neanderthals: What do Fossils, Genetics and Archeology Say? Biolinguistics 7:35–70.Frayer, D., Fiore, I., Lalueza-Fox, C.... & Bondioli, L. (2012). Replay to Benítez-Burraco & Longa: When is enough, enough? Journal of Anthropological Sciences 90:1–6.

abrupt & recentpiecemeal & old

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Some strands of evidence

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Ancient and modern DNA

Admixture Same biological species

Contested hybrid fossils

Dediu, D., & Levinson, S. C. (2013). On the antiquity of language. Frontiers in Language Sciences, 4:397.Disotell, T. R. (2012). Archaic human genomics. American Journal of Physical Anthropology 149:24–39.Currat, M., & Excoffier, L. (2011). Strong reproductive isolation between humans and Neanderthals inferred from observed patterns of introgression. PNAS 108:15129.

- Abrigo do Lagar Vehlo (P)- Peştera cu Oase- Peştera Muierii (RO)- Mladeč (CZ)- Riparo Mezzena (IT)...

- hybridization- multiple definitions of “species”- on the verge of speciation → More interesting question: what processes?

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FOXP2


Lai, C., Fisher, S., Hurst, J., Vargha-Khadem, F., & Monaco, A. (2001). A forkhead-domain gene is mutated in a severe speech and language disorder. Nature 413: 519–23.Spiteri, E., Konopka, G., Coppola, G., Bomar, J., … Geschwind, D. H. (2007). Identification of the Transcriptional Targets of FOXP2, a Gene Linked to Speech and

Language, in Developing Human Brain. The American Journal of Human Genetics 81:1144–1157.

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FOXP2


Fisher, S. E., & Scharff, C. (2009). FOXP2 as a molecular window into speech and language. Trends Genet 25:166–177.Liégeois, F., Morgan, A. T.,... Vargha-Khadem, F. (2011). Endophenotypes of FOXP2: Dysfunction within the human articulatory network. Eur. J. Paed. Neurol. 15:283–288.Vernes, S., Oliver, P., Spiteri, E.,… Lowy, E. (2011). Foxp2 regulates gene networks implicated in neurite outgrowth in the developing brain. PLoS Genetics 7:e1002145.

Multiple phenotypes:

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Fisher, S. E. (2006). Tangled webs: tracing the connections between genes and cognition. Cognition 101:270–297. Fisher, S. E., & Scharff, C. (2009). FOXP2 as a molecular window into speech and language. Trends Genet 25:166–177.

Multiple phenotypes:FOXP2

FOXP2 is not “the” gene “for” language and speech

Simon Fisher

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+ two “human-specific” AAs on same haplotype– intron 8 (POU3F2 binding) → ancestral allele: Africa

FOXP2


Enard, W., Przeworski, M., Fisher, S. E., Lai, C. S. L., … Pääbo, S. (2002). Molecular evolution of FOXP2, a gene involved in speech and language. Nature 418:869–872.Krause, J., Lalueza-Fox, C., … Pääbo, S. (2007). The Derived FOXP2 Variant of Modern Humans Was Shared with Neandertals. Current Biology 17:1908–1912.Maricic, T., Günther, V., … Pääbo, S. (2013). A Recent Evolutionary Change Affects a Regulatory Element in the Human FOXP2 Gene. Molecular Biol. Evol. 30: 844–852.

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+ two “human-specific” AAs on same haplotype– intron 8 (POU3F2 binding) → ancestral allele: Africa

FOXP2

CNTNAP2

MEF2A – developmental plasticity?


Rodenas-Cuadrado, P., Ho, J., & Vernes, S. C. (2013). Shining a light on CNTNAP2: complex functions to complex disorders. European Journal of Human Genetics.Whitehouse, A, Bishop, D, ... Fisher, S (2011). CNTNAP2 variants affect early language development in the general population. Genes, Brain, and Behavior 10:451–456.Somel, M., Liu, X., & Khaitovich, P. (2013). Human brain evolution: transcripts, metabolites and their regulators. Nature Reviews Neuroscience 14:112–127.

- downregulated ↓ by FOXP2- neurexin family- involved in autism, SLI,

normal language variation,language development...

– coding change (Ile → Val) → functional?

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Anatomy & development

Partly due to lifestyle differences

Dediu, D., & Levinson, S. C. (2013). On the antiquity of language. Frontiers in Language Sciences, 4:397.

][

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Birth canalDental

eruption

Prolongedchildhood

MEF2A~0.5mya

Weaver, T. D., & Hublin, J.-J. (2009). Neandertal birth canal shape and the evolution of human childbirth. PNAS 106:8151–8156.Castro, J. M. B. de, Martinón-Torres, M., … Carbonell, E. (2010). New immature hominin fossil from European Lower Pleistocene shows the earliest evidence of a modern

human dental development pattern. PNAS, 107:11739–11744.

][

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Birth canalDental

eruption

Prolongedchildhood

MEF2A~0.5mya

occipitalbunlarger

eyes

comparablebrain size

Cognition?Neural organization?

Klein, R. G. (2009). The Human Career: Human Biological and Cultural Origins (3rd edition). University of Chicago Press. Pearce, E., Stringer, C., & Dunbar, R. I. M. (2013). New insights into differences in brain organization between Neanderthals and anatomically modern humans. Proc. Royal

Soc. B: Biological Sciences 280.

][

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Vocal production & perception

Tuning production ↔ perception

Audiograms

modern human

chimp

Increasedsensitivity

H. heidel bergensis

Kojima, S. (1990). Comparison of auditory functions in the chimpanzee and human. Folia Primatol 55:62–72.Martínez, I., Rosa, M., … Carbonell, E. (2004). Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain. PNAS 101: 9976–9981.Martínez, I., … Arsuaga, J. L. (2013). Communicative capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain. Quaternary International 295:94–101

][[ ]

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Audiograms

Modern humans

Early modern humans

Neandertal

Ear ossicles

Chimp P. robustus

Quam, R., & Rak, Y. (2008). Auditory ossicles from southwest Asian Mousterian sites. Journal of Human Evolution 54:414 – 433. Quam, R. M., Ruiter, D. J. de, Masali, M., Arsuaga, J.-L., Martínez, I., & Moggi-Cecchi, J. (2013). Early hominin auditory ossicles from South Africa. PNAS 110:8847–

8851.

][[ ]

Incus:

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Audiograms

Ear ossicles

Lieberman, P., & Crelin, E. S. (1971). On the speech of Neanderthal man. Linguistic Inquiry 2:203–222.Boer, B. de. (2009). Acoustic analysis of primate air sacs and their effect on vocalization. J Acoust Soc Am 126:3329–3343.

][[ ]

Hyoid bone

air sacks

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Audiograms

Ear ossicles

Hyoid bone

Arensburg, B., Tillier, A. M., Vandermeersch, B., Duday, H., Schepartz, L. A., & Rak, Y. (1989). A Middle Palaeolithic human hyoid bone. Nature 338:758–760.Martínez, I., Arsuaga, J. L., ... Rodríguez, L. (2008). Human hyoid bones from the middle Pleistocene site of the Sima de los Huesos. Journal of Human Evol. 54:118–124.D’Anastasio, R., Wroe, S., … Capasso, L. (2013). Micro-Biomechanics of the Kebara 2 Hyoid and Its Implications for Speech in Neanderthals. PLoS ONE 8:e82261.

][[ ]

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Audiograms

Ear ossicles

Hyoid bone

Breathing control

MacLarnon, A. M., & Hewitt, G. P. (1999). The evolution of human speech: the role of enhanced breathing control. American journal of physical anthropology 109:341–363.

][[ ]

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Audiograms

Ear ossicles

Hyoid bone

Breathing control

All for singing?

Mithen, S. (2005). The Singing Neanderthals: The Origins of Music, Language, Mind and Body. Weidenfeld & Nicolson.

][[ ]

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Symbolic behaviour

Extremely difficult and controversial

Chomsky, N. (2010). Some simple evo devo theses: how true might they be for language. The evolution of human language: biolinguistic perspectives, 62:54–62.Bickerton, D. (2002). From protolanguage to language. In T. Crow (Ed.), The Speciation of Modern Homo Sapiens (pp. 103–120). Oxford: Ofxord University PressDonald, M. (1999). Les origines de l’esprit moderne: Trois étapes dans l’évolution de la culture et de la cognition. DeBoeck Université.

][[ ]

N. Chomsky

I. Tattersall

P. Lieberman

“Modern human revolution” → “modern package”

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Symbolic behaviour


“Modern human revolution”

Chomsky, N. (2010). Some simple evo devo theses: how true might they be for language. The evolution of human language: biolinguistic perspectives, 62:54–62.

][[ ]

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Symbolic behaviour



McBrearty, S., & Brooks, A. S. (2000). The revolution that wasn’t: a new interpretation of the origins of modern human behavior. Journal of Human Evolution 39:453–563.Henshilwood, C. S., & Marean, C. W. (2003). The origin of modern human behavior - Critique of the models and their test implications. Current Anthropology 44:627–651.

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Symbolic behaviour


Complex toolkit (Mousterian): bone, wood, strings...

Thieme, H. (1997). Lower Palaeolithic hunting spears from Germany. Nature 385:807 – 810. Hardy, B. L., Moncel, M.-H., Daujeard, C., … Gallotti, R. (2013). Impossible Neanderthals? Making string, throwing projectiles and catching small game during Marine

Isotope Stage 4 (Abri du Maras, France). Quaternary Science Reviews 82:23–40.


][[ ]

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Symbolic behaviour



Intentional burial, grave offerings

Sick & infirm, medicinal plants

Pettitt, P. B. (2002). The Neanderthal dead: exploring mortuary variability in Middle Palaeolithic Eurasia. Before Farming 1:1–19.Spikins, P. A., Rutherford, H. E., & Needham, A. P. (2010). From Homininity to Humanity: Compassion from the Earliest Archaics to Modern Humans. Time and Mind

3:303–325.


][[ ]

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Symbolic behaviour





Body ornaments: ocher, beads, art

D’ Errico, F., & Soressi, M. (2002). Systematic use of manganese pigment by the Pech-de-l’Azé Neandertals. Journal of Human Evolution 42:A13.Pike, A. W. G., Hoffmann, D. L., … Zilhão, J. (2012). U-Series Dating of Paleolithic Art in 11 Caves in Spain. Science 336:1409–1413.Peresani, M., Vanhaeren, M., ... d’ Errico, F. (2013). An Ochered Fossil Marine Shell From the Mousterian of Fumane Cave, Italy. PLoS ONE 8:e68572.


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Symbolic behaviour






Archaeological absence ⇒ absence of capacity!

Speth, J. D. (2004). News flash: Negative evidence convicts Neanderthals of gross mental incompetence. World Archaeology, 36, 519–526.Henrich, J. (2004). Demography and Cultural Evolution: How Adaptive Cultural Processes can Produce Maladaptive Losses: The Tasmanian Case. American Antiquity

69:197–214.


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Symbolic behaviour






Frayer, D. W., Lozano, M., Bermúdez de Castro, J. M., Carbonell, E., … Bondioli, L. (2012). More than 500,000 years of right-handedness in Europe. Laterality 17:51–69. Volpato, V., Macchiarelli, R., … & Frayer, D. W. (2012). Hand to Mouth in a Neandertal: Right-Handedness in Regourdou 1. PLoS ONE 7:e43949.


][[ ]

Handedness

Archaeological absence ⇒ absence of capacity!

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1Dediu, D., & Levinson, S. C. (2013). On the antiquity of language. Frontiers in Language Sciences, 4:397.Johansson, S. (2013). The Talking Neanderthals: What do Fossils, Genetics and Archeology Say? Biolinguistics 7:35–70.Frayer, D., Fiore, I., Lalueza-Fox, C.... & Bondioli, L. (2012). Replay to Benítez-Burraco & Longa: When is enough, enough? Journal of Anthropological Sciences 90:1–6.

][[ ]

[ ]

][ ?[ ]

[ ] ?

Recognizably modern speech and language (not “proto-language”) at ~0.5mya

Dan Dediu 311

So, how did we get language?

Present~ 2 mya~ 6 mya

MRCA

H. erectus

Neandert(h)als

Denisovans

Chimps

Language

No language

Very probablyno language

Probably language

Probably language

Probably language

Probablyproto-language(s)

Dan Dediu 321

So, how did we get language?

- very many proposals

- confusing literature

- some centuries old

- some (updated versions) are very recent

- Kazuo Okanoya (2010) & Tecumsech Fitch (2012):

- naive evolutionist: continuity, gradualism (e.g., Hockett, Arbib, Tomasello, Pinker,....)

- punctuationist: qualitative differences, saltationism (e.g., Chomsky, Berwick, Tattersall)

- pre-adaptationist: multi-component approach, continuity & innovations, plurality of

mechanisms (e.g, Okanoya, Fitch, Hurford, Leveinson, Dediu,...)

→ exaptationist: “[...] cascade of innovations, each creating the preconditions for

the latter ones to be functionally and adaptively favored” (Fitch 2012:631)

Okanoya, K. (2010). Biological pre-adaptation for language emergence. Kyoto, Japan: EELC 2010 Emergence and Evolution of Linguistic Communication.Fitch, W. T. (2012). Evolutionary Developmental Biology and Human Language Evolution: Constraints on Adaptation. Evolutionary Biology 39:613–637.

Dan Dediu 331

Problem:everybody claims to be “Evo-Devo”

- famous saltationists have recently argued that Evo-Devo supports their claims:

- Chomsky (2010):

- Benítez-Burraco & Longa (2010): argue that Chomsky's saltationism (in what concerns

minimalism) is supported by evo-devo, just not the version that Chomsky himself has

picked (“Evo-DevoGEN

” aka Sean Carroll's), but by DST/phenotypic plasticity)

- interestingly, Benítez-Burraco & Boeckx (2014) [Universal Grammar and Biological Variation: An EvoDevo

Agenda for Comparative Biolinguistics. Biological Theory 9:122–134] apparently argue that widespread variation

goes against Universal Grammar and instead Evo-Devo concepts such as developmental

plasticity, robustness/canalization and evolvability are better suited

Chomsky, N. (2010). Some simple evo devo theses: how true might they be for language. The evolution of human language: biolinguistic perspectives, 62:54–62.Berwick, R., & Chomsky, N. (2011). The biolinguistic program: The current state of its development. In A. M. di Sciullo & C. Boeckx (Eds.), OUPBenítez-Burraco, A., & Longa, V. M. (2010). Evo-devo—of course, but which one? Biolinguistics 4:308–323.

“a rewiring of the brain took place in some individual, call him Prometheus, yielding the operation of unbounded Merge, applying to concepts with intricate (and little understood) properties…”

Dan Dediu 341

Problem:everybody claims to be “Evo-Devo”

- but some go even further: Fodor & Piatelli-Palmarini (2010) criticize natural selection

→ Douglas Futuyma (2010) famously wrote:

“Because they are prominent in their own fields, some readers may suppose that they are authorities

on evolution who have written a profound and important book. They aren't, and it isn't.”

- Bickerton (2014):

“[...] the attitudes of many biolinguists towards natural selection and evo-devo: The first they

misunderstand, the second they both misunderstand and overestimate.”

Fodor, J., & Piattelli-Palmarini, M. (2010). What Darwin Got Wrong. Farrar, Straus and Giroux.Futuyma, D. J. (2010). Two Critics Without a Clue. Science 328:692–693Bickerton, D. (2014). Some problems for biolinguistics. Biolinguistics 8:073–096.

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So, where's the truth?

1.) we need to move beyond speculations and towards actual empirical findings:

- child language acquisition, computational modeling, genetics, comparative work...

2.) almost trivially an evo-devo approach to language is part of the solution:

- however, the theory must fit the data not the other way around

- and how far can we push the metaphors?

3.) but language (and culture) need more than evo-devo!

→

Carroll, S. B. (2011). Endless Forms Most Beautiful: The New Science of Evo Devo : and the Making of the Animal Kingdom. Quercus Publishing. West-Eberhard, M. J. (2003). Developmental Plasticity and Evolution (1st ed.). NY: Oxford University Press.

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Language (and culture)need more than Evo-Devo

1.) it is probably wrong to qualitatively distinguish language evolution from language change

→ linguistic diversity and universal tendencies are both essential phenomena

→ language is a full-fledged evolutionary system in its own:

- phylogenetic/phylogeographic methods

- rates of evolution, constrained & biased evolution....

- specific phenomena: massive reticulation, directed change

Richerson, P. J., & Christiansen, M. (Eds.). (2013). Cultural evolution: society, technology, language, and religion (Vol. 12). Cambridge, Mass: MIT Press.Atkinson, Q. D., & Gray, R. D. (2005). Curious parallels, curious connections - Phylogenetic Thinking in Biology and Historical Linguistics. Systematic Biology 54:513-526.Bouckaert, R., Lemey, P., Dunn, M.,… Atkinson, Q. D. (2012). Mapping the Origins and Expansion of the Indo-European Language Family. Science 337: 957–960.

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Language (and culture)need more than Evo-Devo

2.) language (and culture) are active and powerful evolutionary forces:

→ Mayr's proximate ↔ ultimate distinction is blurred (Laland et al. 2011)

→ pervasive (cultural) niche construction (Odling-Smee et al. 2003)

→ gene-culture co-evolution & biased cultural evolution (Levinson & Dediu, 2013)

- e.g., vocal tract ↔ speech co-evolution

⇒ (eco-)evo-devo is a foundation on which to build but is not the complete model!

Laland, K.., Sterelny, K., et al.. (2011). Cause and Effect in Biology Revisited: Is Mayr’s Proximate-Ultimate Dichotomy Still Useful? Science 334:1512-1516.Odling-Smee, F. J., Laland, K. N., & Feldman, M. W. (2003). Niche Construction: The Neglected Process in Evolution. Princeton University Press.Levinson, S. C., & Dediu, D. (2013). The interplay of genetic and cultural factors in ongoing language evolution. In Richerson, P. & Christiansen, M. MIT Press.

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Conclusions

Thank you!Acknowledgments: Kim Oller, Ulrike Griebel, Steve Levinson, Sonja Vernes

& the Language and Genetics department

Funded by Netherlands Organisation for Scientific Research ( ) VIDI grant 276-70-022

Language is old...

… and requires (more than) evo-devo

modern evolutionary thinking in language evolution and … · modern evolutionary thinking in...

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