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Dan Dediu 1
LSA2013Universality and Variability:
New Insights from Genetics29th of June, 2013
Dan Dediu
Language and GeneticsMax Planck Institute for Psycholinguistics
NijmegenThe Netherlands
1
Genes and culture inlanguage evolution
Dan Dediu 21Overview
● Population and evolutionary genetics
● Human evolution
● Human genetic and linguistic diversity
● Genetic biasing, gene-culture co-evolution, niche construction
Overview
Dan Dediu 31Population and evolutionary genetics
Population and evolutionary genetics
cell membrane
mitochondria nucleus
nuclearmembrane
mitochondrialmembrane
mitochondrialDNA (mtDNA)
nuclearDNA
Dan Dediu 41Population and evolutionary genetics
Population and evolutionary genetics
cell membrane
mitochondria nucleus
nuclearmembrane
mitochondrialmembrane
mitochondrialDNA (mtDNA)
nuclearDNA
Genome
Dan Dediu 51Population and evolutionary genetics
Population and evolutionary genetics
● Some genes (energy production)
● Own genetic code● “D-loop” → hypervariable
regions (HVR-I & HVR-II)
Dan Dediu 61Population and evolutionary genetics
Population and evolutionary genetics
Maternal inheritance
● Some genes (energy production)
● Own genetic code● “D-loop” → hypervariable
regions (HVR-I & HVR-II)
Dan Dediu 71Population and evolutionary genetics
Population and evolutionary genetics
● Chromosomes: single linear molecules of DNA● Karyotype = all chromosomes in the nucleus● Humans: 23 pairs of chromosomes (diploid)
● 22 pairs autosomes● 1 pair sex chromosomes
● XX=♀ XY=♂● Inheritance:
● X & autosomes: both parents
● Y: paternal inheritance
Dan Dediu 81Population and evolutionary genetics
Population and evolutionary genetics
X Y♂
X X♀
autosome pair
sex chromosomes
mtDNA
X Y♂
X Y♂
X X♀
parents
offspring
● the inheritance of the nuclear genome● recombination, independent assortment
Dan Dediu 91Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
AaAA
aa
Aa
AAaa
AaAa
Aa
Population
Dan Dediu 101Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
AaAA
aa
Aa
AAaa
AaAa
Aa
IndividualsPopulation
Dan Dediu 111Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
AaAA
aa
Aa
AAaa
AaAa
Aa
Individuals
HeterozygousHomozygous
Population
Dan Dediu 121Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
● Allele frequencies pA & p
a
● Genotype frequencies pAA
, pAa
& paa
AaAA
aa
Aa
AAaa
AaAa
Aa
Population
Dan Dediu 131Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
● Allele frequencies pA & p
a
● Genotype frequencies pAA
, pAa
& paa
AaAA
aa
Aa
AAaa
AaAa
Aa
Time
tp
A, p
a; p
AA, p
Aa & p
aa
Dan Dediu 141Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
● Allele frequencies pA & p
a
● Genotype frequencies pAA
, pAa
& paa
AaAA
aa
Aa
AAaa
AaAa
AaAa
AA
aa
Aa
AA
aa
AaAa
Aa
Time
t t+1
Aa
aa
aa
Aa aa
aa
AA
pA, p
a; p
AA, p
Aa & p
aap'
A, p'
a; p'
AA, p'
Aa & p'
aa
Dan Dediu 151Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
● Allele frequencies pA & p
a
● Genotype frequencies pAA
, pAa
& paa
AaAA
aa
Aa
AAaa
AaAa
AaAa
AA
aa
Aa
AA
aa
AaAa
Aa
Time
t t+1
Aa
aa
aa
Aa aa
aa
AA
pA, p
a; p
AA, p
Aa & p
aap'
A, p'
a; p'
AA, p'
Aa & p'
aa
conditions &factors
Dan Dediu 161Population and evolutionary genetics
Population and evolutionary genetics
● Bi-allelic autosomal locus → A & a
● Allele frequencies pA & p
a
● Genotype frequencies pAA
, pAa
& paa
AaAA
aa
Aa
AAaa
AaAa
AaAa
AA
aa
Aa
AA
aa
AaAa
Aa
Time
t t+1
Aa
aa
aa
Aa aa
aa
AA
pA, p
a; p
AA, p
Aa & p
aap'
A, p'
a; p'
AA, p'
Aa & p'
aa
conditions &factors
...
t+n
Dan Dediu 171Population and evolutionary genetics
Population and evolutionary genetics
● Hardy-Weinberg Equilibrium (HWE):
if nothing interesting happens → no change
AaAA
aa
Aa
AAaa
AaAa
Aa
Time
tp
A, p
a; p
AA, p
Aa & p
aa
Dan Dediu 181Population and evolutionary genetics
Population and evolutionary genetics
● Hardy-Weinberg Equilibrium (HWE):
if nothing interesting happens → no change
AaAA
aa
Aa
AAaa
AaAa
Aa
Time
tp
A, p
a; p
AA, p
Aa & p
aa
● infinite population● no mutation● no natural selection● random mating● no migration
AaAA
aa
Aa
AAaa
AaAa
Aa
pA, p
a; p
AA, p
Aa & p
aa
t+1
Dan Dediu 191Population and evolutionary genetics
Population and evolutionary genetics
● Hardy-Weinberg Equilibrium (HWE):
if nothing interesting happens → no change
AaAA
aa
Aa
AAaa
AaAa
Aa
Time
tp
A, p
a; p
AA, p
Aa & p
aa
● infinite population● no mutation● no natural selection● random mating● no migration
AaAA
aa
Aa
AAaa
AaAa
Aa
pA, p
a; p
AA, p
Aa & p
aa
t+1
Dan Dediu 201Population and evolutionary genetics
Population and evolutionary genetics
● Hardy-Weinberg Equilibrium (HWE):
if nothing interesting happens → no change
→ simple null hypothesis
AaAA
aa
Aa
AAaa
AaAa
Aa
Time
tp
A, p
a; p
AA, p
Aa & p
aa
● infinite population● no mutation● no natural selection● random mating● no migration
AaAA
aa
Aa
AAaa
AaAa
Aa
t+1p
A, p
a; p
AA, p
Aa & p
aa
Dan Dediu 211Population and evolutionary genetics
Population and evolutionary genetics
● Finite populations → reproduction = random sampling
Dan Dediu 221Population and evolutionary genetics
Population and evolutionary genetics
● Finite populations → reproduction = random sampling
● Bottlenecks → “founder effect” (e.g., SLI on Robinson Crusoe Island)
Dan Dediu 231Population and evolutionary genetics
Population and evolutionary genetics
● Finite populations → reproduction = random sampling
● Bottlenecks → “founder effect”
● (Effective) population size, Ne
Dan Dediu 241Population and evolutionary genetics
Population and evolutionary genetics
● Finite populations → reproduction = random sampling
● Bottlenecks → “founder effect”
● (Effective) population size, Ne
→ reduces genetic diversity
Dan Dediu 251Population and evolutionary genetics
Population and evolutionary genetics
● Mutation → changes the genetic information (e.g., A → a)
● Many types: point mutation
Dan Dediu 261Population and evolutionary genetics
Population and evolutionary genetics
● Mutation → changes the genetic information (e.g., A → a)
● Many types: point mutation, chromosomal rearrangements
Dan Dediu 271Population and evolutionary genetics
Population and evolutionary genetics
● Mutation → changes the genetic information (e.g., A → a)
● Many types: point mutation, chromosomal rearrangements, gene duplications
Dan Dediu 281Population and evolutionary genetics
Population and evolutionary genetics
● Mutation → changes the genetic information (e.g., A → a)
● Many types: point mutation, chromosomal rearrangements, gene duplications, CNVs...
Dan Dediu 291Population and evolutionary genetics
Population and evolutionary genetics
● Mutation → changes the genetic information (e.g., A → a)
● Many types: point mutation, chromosomal rearrangements, gene duplications, CNVs...
→ increases genetic diversity
Dan Dediu 301Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
Dan Dediu 311Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
natural
Dan Dediu 321Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
natural sexualvs
Dan Dediu 331Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
natural sexual artificialvs vs
Dan Dediu 341Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
vs vs
Dan Dediu 351Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
vs vs
Stabilizing
Dan Dediu 361Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
vs vs
Stabilizing Directional
Dan Dediu 371Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
vs vs
Stabilizing Directional Disruptive
Dan Dediu 381Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Types:
vs vs
Stabilizing Directional Disruptive Heterozygoteadvantage
Dan Dediu 391Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Walking the fitness landscape
Dan Dediu 401Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Walking the fitness landscape
Dynamic fitness landscape
Dan Dediu 411Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Walking the fitness landscape
Co-evolution & arms racesDynamic fitness landscape
Dan Dediu 421Population and evolutionary genetics
Population and evolutionary genetics
● Selection → differential survival and reproduction
● Walking the fitness landscape
Co-evolution & arms races
Niche construction
Dynamic fitness landscape
Dan Dediu 431Population and evolutionary genetics
Population and evolutionary genetics
● Non-random mating
Dan Dediu 441Population and evolutionary genetics
Population and evolutionary genetics
● Non-random mating
– inbreeding
Dan Dediu 451Population and evolutionary genetics
Population and evolutionary genetics
● Non-random mating
– inbreeding
– assortative mating
Dan Dediu 461Population and evolutionary genetics
Population and evolutionary genetics
● Non-random mating
– inbreeding
– assortative mating
– population subdivision
Dan Dediu 471Population and evolutionary genetics
Population and evolutionary genetics
● Non-random mating
– inbreeding
– assortative mating
– population subdivision
→ gene flow/admixture
Dan Dediu 481Human evolution
Human evolution
Dan Dediu 491Human evolution
Human evolution
Tree of life
Dan Dediu 501Human evolution
Human evolution
Tree of life
Dan Dediu 511Human evolution
Human evolution
Tree of life
Dan Dediu 521Human evolution
Human evolution
Dan Dediu 531Human evolution
Human evolution
Dan Dediu 541Human evolution
Human evolution
Dan Dediu 551Human evolution
Human evolution
Dan Dediu 561Human evolution
Human evolution
Dan Dediu 571Human evolution
Human evolution
Dan Dediu 581Human evolution
Human evolution
Dan Dediu 591Human evolution
Human evolution
Dan Dediu 601Human evolution
Human evolution
Dan Dediu 611Human evolution
Human evolution
1-4% (2.5%) non-Africans
~4% Melanesians
Dan Dediu 621Human evolution
Human evolution
● “Modern” exon FOXP2
● Hyoid bone
● Audiograms
● Tools
● Symbolic culture (?)
=
Dan Dediu 631Human evolution
Human evolution
● “Modern” exon FOXP2
● Hyoid bone
● Audiograms
● Tools
● Symbolic culture (?)
=
● Intron 8 FOXP2/POU3F2
● CNTNAP2
● MCPH1, ASPM, DRD5, MEF2A
● Survival
● Symbolic culture, tools...
≠
Dan Dediu 641Human evolution
Human evolution
● “Modern” exon FOXP2
● Hyoid bone
● Audiograms
● Tools
● Symbolic culture (?)
=
● Intron 8 FOXP2/POU3F2
● CNTNAP2
● MCPH1, ASPM, DRD5, MEF2A
● Survival
● Symbolic culture, tools...
≠
Did “they” have language?If so, what kind?
Dan Dediu 651Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Genetically quite similar to other animals:
– 98% to chimps
Dan Dediu 661Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Genetically quite similar to other animals:
– 98% to chimps
– 85% to mice
98%98%
Dan Dediu 671Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Genetically quite similar to other animals:
– 98% to chimps
– 85% to mice
– 50% to fruit flies
98%98% 85%85%
Dan Dediu 681Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Genetically quite similar to other animals:
– 98% to chimps
– 85% to mice
– 50% to fruit flies● Most differences in regulatory networks and dynamics
98%98% 85%85% 50%50%
Dan Dediu 691Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
Dan Dediu 701Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
~ <0.5% nucleotide diversity b/w individuals (15 million bases)
Dan Dediu 711Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
Dan Dediu 721Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
Dan Dediu 731Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
Dan Dediu 741Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
diversity decreases with distance from Africa
Dan Dediu 751Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
diversity decreases with distance from Africa
phenotype
Dan Dediu 761Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
diversity decreases with distance from Africa
phenotype genotype
Dan Dediu 771Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
diversity decreases with distance from Africa
gradual clines as opposed to discontinuous boundaries
Dan Dediu 781Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Relatively genetically uniform species ← evolutionary history
● But we are no clones!
● Distribution of diversity
~ 8% between : 7% within continents : 85% within local groups
diversity decreases with distance from Africa
gradual clines as opposed to discontinuous boundaries
due to ubiquitous allele frequencies, less to private alleles:
Dan Dediu 791Human genetic and linguistic diversity
Human genetic and linguistic diversity
– continuous
Distribution of Y haplogroups (Jobling & Tyler-Smith 2003)
Dan Dediu 801Human genetic and linguistic diversity
Human genetic and linguistic diversity
Distribution of Y haplogroups (Jobling & Tyler-Smith 2003)
Human genetic diversity(Barbujani & Colonna, 2010)
– continuous
– multiple loci
Dan Dediu 811Human genetic and linguistic diversity
Human genetic and linguistic diversity
Distribution of Y haplogroups (Jobling & Tyler-Smith 2003)Genetic diversity Europe (500,000 SNPs)
Human genetic diversity(Barbujani & Colonna, 2010)
– continuous
– multiple loci
– geography
Dan Dediu 821Human genetic and linguistic diversity
Human genetic and linguistic diversity
● ~ 7000 languages● > 100 language families
● ~ 150 incl. language isolates
Dan Dediu 831Human genetic and linguistic diversity
Human genetic and linguistic diversity
● ~ 7000 languages● > 100 language families
● ~ 150 incl. language isolates
IE tree with ages, Gray & Atkinson (2003)
● Language = cultural evolutionary system● vertical inheritance → language families● horizontal processes → language areas
Indian sprachbund
Dan Dediu 841Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Humans carry genes & languages → historical processes should create or break correlations
● Migrations
Viking longboat
Dan Dediu 851Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Humans carry genes & languages → historical processes should create or break correlations
● Migrations● Conquest
Roman legion
Dan Dediu 861Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Humans carry genes & languages → historical processes should create or break correlations
● Migrations● Conquest● Religion
Sultan Ahmed Mosque, Istambul
Dan Dediu 871Human genetic and linguistic diversity
Human genetic and linguistic diversity
● Farming/language co-dispersal
Proposed farming expansions Diamond & Bellwood (2003)
Dan Dediu 881Human genetic and linguistic diversity
Human genetic and linguistic diversity
→ accidental correlations
Dan Dediu 891Human genetic and linguistic diversity
Human genetic and linguistic diversity
→ accidental correlations→ non-accidental correlations:
Dan Dediu 901Human genetic and linguistic diversity
Human genetic and linguistic diversity
→ accidental correlations→ non-accidental correlations:● village sign languages
Kata Kolok
ABSL
Dan Dediu 911Human genetic and linguistic diversity
Human genetic and linguistic diversity
→ accidental correlations→ non-accidental correlations:● village sign languages● tone, ASPM, Microcephalin
ASPM
Mic
roce
phal
in
http://wals.info
Dan Dediu 921Human genetic and linguistic diversity
Human genetic and linguistic diversity
→ accidental correlations→ non-accidental correlations:● village sign languages● tone, ASPM, Microcephalin → genetic biasing
Dan Dediu 931Niche construction & gene-culture co-evolution
Niche construction &Gene-culture co-evolution
● Organisms construct their niches
Niche construction
Selective pressures
Dan Dediu 941Niche construction & gene-culture co-evolution
● Organisms construct their niches
● Culture shapes selective pressures:
– farming
Niche construction &Gene-culture co-evolution
Dan Dediu 951Niche construction & gene-culture co-evolution
● Organisms construct their niches
● Culture shapes selective pressures:
– farming → immune system
Niche construction &Gene-culture co-evolution
Dan Dediu 961Niche construction & gene-culture co-evolution
● Organisms construct their niches
● Culture shapes selective pressures:
– farming → immune system, amylase gene
Niche construction &Gene-culture co-evolution
Dan Dediu 971Niche construction & gene-culture co-evolution
● Organisms construct their niches
● Culture shapes selective pressures:
– farming → immune system, amylase gene, lactose tolerance...
Niche construction &Gene-culture co-evolution
Dan Dediu 981Niche construction & gene-culture co-evolution
● Organisms construct their niches
● Culture shapes selective pressures:
– farming → immune system, amylase gene, lactose tolerance...
– tool use → hand
Niche construction &Gene-culture co-evolution
Dan Dediu 991Niche construction & gene-culture co-evolution
● Organisms construct their niches
● Culture shapes selective pressures:
– farming → immune system, amylase gene, lactose tolerance...
– tool use → hand
– speech → vocal tract
Niche construction &Gene-culture co-evolution
Dan Dediu 100
1Niche construction & gene-culture co-evolution
● Organisms construct their niches
● Culture shapes selective pressures:
– farming → immune system, amylase gene, lactose tolerance...
– tool use → hand
– speech → vocal tract
– language → whole world
Niche construction &Gene-culture co-evolution
Dan Dediu 101
1Niche construction & gene-culture co-evolution
Conclusions
● Human evolution was really complicated → no simple story
● Complex relationships between genes and language
→ co-evolution and niche construction
● Ongoing evolution of the genetic architecture for language?
Dan Dediu 102
1
Conclusions
● Human evolution was really complicated → no simple story
● Complex relationships between genes and language
→ co-evolution and niche construction
● Ongoing evolution of the genetic architecture for language?
Thanks to: Alejandrina Cristia, Sarah Graham, Steve Levinson
Funding: Netherlands Organisation for Scientific Research ( ) Vidi grant 276-70-022
Dan Dediu 103
1
Suggested reading
Suggested readings
● Books:Jobling, M. A., Hurles, M., & Tyler-Smith, C. (2004). Human Evolutionary Genetics: Origins, Peoples and Disease. Garland Science: NY.
Halliburton, R. (2004). Introduction to population genetics. Pearson Eduction Inc.: Upper Saddle River.
● Population and evolutionary genetics:Hurst, L. D. (2009). Fundamental concepts in genetics: genetics and the understanding of
selection. Nat Rev Genet 10:83–93. doi:10.1038/nrg2506
Charlesworth, B. (2009). Fundamental concepts in genetics: effective population size and patterns of molecular evolution and variation. Nat Rev Genet 10:195–205. doi:10.1038/nrg2526
Jablonka, E., & Raz, G. (2009). Transgenerational epigenetic inheritance: prevalence, mechanisms, and implications for the study of heredity and evolution. The Quarterly Review of Biology 84:131–176.
Koonin, E. V. (2009). Darwinian evolution in the light of genomics. Nucleic Acids Res 37:1011–1034. doi:10.1093/nar/gkp089
Dan Dediu 104
1
Suggested reading
Suggested readings
● Human evolution:Green, R. E., Krause, J., Briggs, A. W., Maricic, T., Stenzel, U., Kircher, M., … Fritz, M.
H.-Y. (2010). A draft sequence of the Neandertal genome. Science 328:710–722. doi:10.1126/science.1188021
Disotell, T. R. (2012). Archaic human genomics. American Journal of Physical Anthropology 149:24–39. doi:10.1002/ajpa.22159
Meyer, M., Kircher, M., Gansauge, M.-T., Li, H., Racimo, F., Mallick, S., … Pääbo, S. (2012). A High-Coverage Genome Sequence from an Archaic Denisovan Individual. Science 338:222–226. doi:10.1126/science.1224344
Dediu, D., & Levinson, S. C. (2013). On the antiquity of language: the reinterpretation of Neandertal linguistic capacities and its consequences. Frontiers in Language Sciences 4:397. doi:10.3389/fpsyg.2013.00397
● Human diversity:Barbujani, G., & Colonna, V. (2010). Human genome diversity: frequently asked
questions. Trends in Genetics 26:285–295. doi:10.1016/j.tig.2010.04.002
Novembre, J., Johnson, T., Bryc, K., Kutalik, Z., Boyko, A. R., Auton, A., … Bustamante, C. D. (2008). Genes mirror geography within Europe. Nature 456:98–101. doi:10.1038/nature07331
Dan Dediu 105
1
Suggested reading
Suggested readings
● Linguistic and genetic diversity:Diamond, J., & Bellwood, P. (2003). Farmers and their languages: the first expansions.
Science 300:597–603.
Dediu, D. (2011). Are languages really independent from genes? If not, what would a genetic bias affecting language diversity look like? Hum Biol 83:279–296. doi:10.3378/027.083.0208
● Gene-culture co-evolution/niche construction:Laland, K. N., Odling-Smee, J., & Myles, S. (2010). How culture shaped the human
genome: bringing genetics and the human sciences together. Nat Rev Genet 11:137–148. doi:10.1038/nrg2734
Fisher, S. E., & Ridley, M. (2013). Culture, Genes, and the Human Revolution. Science 340:929–930. doi:10.1126/science.1236171
Senghas, A. (2005). Language emergence: clues from a new bedouin sign. Curr Biol 15:R463–R465. doi:10.1016/j.cub.2005.06.018
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