Book: The Agile Gene

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Book 1, Line 58.

Observer, 11 Feb. 2001.

San Francisco Chronicle, 11 Feb. 2001.

New York Times, 12 Feb. 2001.



Act 3, scene 4.

Keynes, R.D. (ed.). 1988. Charles Darwin’s Beagle Diary. Cambridge University Press.


Keynes, R.D. 2001. Annie’s Box. 4th Estate.

Quoted in Degler, C.N. 1991. In Search of Human Nature. Oxford University Press.

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Call, J. 2001. Chimpanzee social cognition. Trends in Cognitive Science 5:388–93.

Malik, K. 2001. What Is It to Be Human? Institute of Ideas.

Darwin, C. 1871. The Descent of Man. John Murray.

Malik, K. 2001. What Is It to Be Human? Institute of Ideas.

Midgley, M. 1978. Beast and Man. Routledge.

Zuk, M. 2002. Sexual Selections. University of California Press.

van Schaik, C.P. and Kappeler, P.M. 1997. Infanticide risk and the evolution of male–female association in primates. Proceedings of the Royal Society B:264:1687–94.

Wrangham, R.W., Jones, J.H., Laden, G., Pilbeam, D., and Conkin-Brittain, N. 1999. The raw and the stolen. Cooking and the ecology of human origins. Current Anthropology 40:567–94.

Ridley, M. 1996. The Origins of Virtue. Penguin.

Wrangham, R.W. and Peterson, D. 1997. Demonic Males. Bloomsbury.

Alan Dixson, e-mail correspondence.

Ebersberger, I., Metzier, D., Schwarz, C., and Paabo, S. 2002. Genome-wide comparison of DNA sequences between human and chimpanzees. American Journal of Human Genetics 70:1490–97.

Britten, R.J. 2002. Divergence between samples of chimpanzee and human DNA sequences is 5%, counting indels. Proceedings of the National Academy of Sciences 99:13633–5.

King, M.C. and Wilson, A.C. 1975. Evolution at two levels in humans and chimpanzees. Science 188:107–16.

Sibley, C.G. and Ahiquist, J.E. 1984. The phylogeny of the hominoid primates, as indicated by DNA-DNA hybridization. Journal of Molecular Evolution 20:2–15.

Johnson, M.E., Viggiano, L., Bailey, J.A., Abdul-Rauf, M., Goodwin, G., Rocchi, M., and Eichler, E.E. 2001. Positive selection of a gene during the emergence of humans and African apes. Nature 413:514–19.

Hayakawa, T., Satta, Y., Gagneux, P., Varki, A., and Takahata, N. 2001. Alu-mediated inactivation of the human CMP-N-acetyineuraminic acid hydroxylase gene. Proceedings of the National Academy of Sciences 98:11399–404.

Ajit Varki, interview. See also Chou, H.-H. et al. 1998. A mutation in human CMP-sialic acid hydroxylase occurred after the Homo-Pan divergence. Proceedings of the National Academy of Sciences 95:11751–6; Gagneux, P. and Varki, A. 2001. Genetic differences between humans and great apes. Molecular Phylogenetics and Evolution 18:2–13; Varki, A. 2001. Loss of N-glycolylneuraminic acid in humans: Mechanisms, consequences, and implications for hominid evolution. Yearbook of Physical Anthropology 44:54–69

Hammer, C.J., Tyler, H.D., Loskutoff, N.M., Armstrong, D.L., Funk, D.J., Lindsey, B.R., and Simmons, L.G. 2001. Compromised development of calves (Bos gaurus) derived from in vitro-generated embryos and transferred interspecifically into domestic cattle (Bos taurus). Theriogenology 55:1447–55; Loskutoff, N., e-mail correspondence.

There is some confusion over terminology here. Some biologists use “promoter” to mean the site where the RNA polymerase enzyme binds after being recruited by a transcription factor. Here I use it in the broader sense, to mean the entire regulatory sequence of the gene.

Belting, H.G., Shashikant, C.S., and Ruddle, F.H. 1998. Modification of expression and cis-regulation of Hoxc8 in the evolution of diverged axial morphology. Proceedings of the National Academy of Sciences 95:2355–60.

Cohn, M.J. and Tickle, C. 1999. Developmental basis of limblessness and axial patterning in snakes. Nature 399:474–9.

Ptashne, M. and Gann, A. 2002. Genes and Signals. Cold Spring Harbor Press; also Alex Gann, interviews.

Carroll, S.B. 2000. Endless forms: The evolution of gene regulation and morphological diversity. Cell 101:577–80.

Coppinger, R. and Coppinger, L. 2001. Dogs: A Startling New Understanding of Canine Origin, Behavior, and Evolution. Scribner.

Semendeferi, K., Armstrong, E, Schleicher, A., Zilles, K., and van Hoesen, G. W. 1998. Limbic frontal cortex in hominoids: A comparative study of area 13. American Journal of Physical Anthropology 106:129–55.

Wrangham, R.W., Pilbeam, D., and Hare, B. (Unpublished.) Convergent paedomorphism in bonobos, domesticated animals, and humans: The role of selection for reduced aggression.

Wrangham, R.W. and Pilbeam, D. 2001, in All Apes Great and Small, volume 1, Chimpanzees, Bonobos, and Gorillas (ed. Galdikas, B., Erickson, N., and Sheeran, L.K.). Plenum; also Wrangham, R.W. Talk at Cold Spring Harbor, President’s Council, May 2001.

Quoted in the New York Times, 24 September 2002.

Bond, J., Roberts, E., Mochida, G.H., Hampshire, D.J., Scott, S., Askham, J.M., Springell, K., Mahadevan, M., Crow, Y.J., Markham, A.F., Walsh, C.A., and Woods, C.G. 2002. ASPM is a major determinant of cerebral cortical size. Nature Genetics 32:316–20.


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Myers, G.E. 1986. William James: His Life and Thought. Yale University Press.

Bender, B. 1996. The Descent of Love: Darwin and the Theory of Sexual Selection in American Fiction, 1871–1926. University of Pennsylvania Press.

James, W. 1890. The Principles of Psychology. Henry Holt.

Myers, G.E. 1986. William James: His Life and Thought. Yale University Press.

Dawkins, R. 1986. The Blind Watchmaker. Norton.

Dennett, D. 1995. Darwin’s Dangerous Idea. Penguin.

James, W. 1890. The Principles of Psychology. Henry Holt.

Insel, T.R. and Shapiro, L.E. 1992. Oxytocin receptor distribution reflects social organization in monogamous and polygamous voles. Proceedings of the National Academy of Sciences 89:5981–5.

Argiolas, A., Melis, M.R., Stancampiano, R., and Gessa, G.L. 1989. Penile erection and yawning induced by oxytocin and related peptides: Structure-activity relationship. Peptides 10:559–63.

Insel, T.R. and Shapiro, L.E. 1992. Oxytocin receptor distribution reflects social organization in monogamous and polygamous voles. Proceedings of the National Academy of Sciences 89:5981–5.

Ferguson, J.N., Young, L.J., Hearn, E.F., Matzuk, M.M., Insel, T.R., and Winslow, J.T. 2000. Social amnesia in mice lacking the oxytocin gene. Nature Genetics 25:284–8.

Young, L.J., Wang, Z., and Insel, T.R. 1998. Neuroendocrine bases of monogamy. Trends in Neurosciences 21:71–5.

Insel, T.R., Winslow, J.T., Wang, Z., and Young, L.J. 1998. Oxytocin, vasopressin, and the neuroendocrine basis of pair bond formation. Advances in Experimental and Medical Biology 449:215–24.

Insel, T.R. and Young, L.J. 2001. The neurobiology of attachment. Nature Reviews in Neuroscience 2:129–36.

Wang, Z., Yu, G., Cascio, C., Liu, Y., Gingrich, B., and Insel, T.R. 1999. Dopamine D2 receptor-mediated regulation of partner preference in female prairie voles (Microtus ochrogaster): A mechanism for pair bonding? Behavioral Neuroscience 113:602–11.

Jankowiak, W.R. and Fisher, E.F. 1992. A cross-cultural perspective on romantic love. Ethnology 31:149–55.

Insel. T.R., Gingrich, B.S., and Young, L.J. 2001. Oxytocin: Who needs it? Progress in Brain Research 133:59–66.

Bartels, A. and Zeki, S. 2000. The neural basis of romantic love. Neuroreport 11:3829–34.

Carter, C.S. 1998. Neuroendocrine perspectives on social attachment and love. Psychoneuroendocrinology 23:779–818.

Ridley, M. 1993. The Red Queen. Penguin.

Tinbergen, N. 1951. The Study of Instinct. Oxford University Press.

Ginsburg, B.E. 2001. Fellow travellers on the road to the genetics of behavior: Mice, rats, and dogs. Talk to the International Behavioural and Neural Genetics Society, 8–10 November 2001, San Diego.

Konner, M. 2001. The Tangled Wing: Biological Constraints on the Human Spirit. 2nd edition. W.H. Freeman.

Budiansky, S. 2000. The Truth about Dogs. Viking Penguin.

You can find such a bull catalog at

Eib-Eibesfeldt, I. 1989. Human Ethology. Aldine de Gruyter; Ekman, P. 1998. Afterword: Universality of emotional expression? A personal history of the dispute. In Darwin, C., The Expression of the Emotions in Man and Animals (new edition). Oxford University Press.

Buss, D.M. 1994. The Evolution of Desire. Basic Books.

Buss, D.M. 2000. The Dangerous Passion. Bloomsbury.

You can find this quoted almost anywhere on the Internet.

Diamond, M., 1965. A critical evaluation of the ontogeny of human sexual behavior. Quarterly Review of Biology 40:147–75.

Colapinto, J. 2000. As Nature Made Him: The Boy Who Was Raised as a Girl. HarperCollins.

Reiner, W.G. 1999. Assignment of sex in neonates with ambiguous genitalia. Current Opinion in Pediatrics 11:363–5. Also article in the Times (London), 26 June 2001, by Lisa Melton: Ethics and gender.

Lutchmaya, S., Baron-Cohen, S., and Raggatt, P. In press. Foetal testosterone and eye contact in 12 month old human infants. Infant Behavior Development.

Connellan, H., Baron-Cohen, S., Wheelwright, S., Batki, A., and Ahluwalia, J. 2000. Sex differences in human neonatal social perception. Infant Behavior and Development 23:113–18.

Baron-Cohen, S. 2002. The extreme male brain theory of autism. Trends in Cognitive Sciences 6:248–54.

Baron-Cohen, S. 2002. Autism: Deficits in folk psychology exist alongside superiority in folk physics. In Understanding Other Minds (ed. Baron-Cohen, S., Tager-Flusberg, H., and Cohen, D.J.), pp. 73–82; Baron-Cohen, S., Wheelwright, S., Skinner, R., Martin, J., and Clubley, E. 2001. The autism spectrum quotient: Evidence from Asperger syndrome high-functioning autism, males and females, scientists and mathematicians. Journal of Autism and Developmental Disorders 31:5–17.

Baron-Cohen, S., interview.

Frith, C. and Frith, U. 2000. The physiological basis of theory of mind: Functional neuroimaging studies. In Understanding Other Minds (ed. Baron-Cohen, S., Tager-Flusberg, H., and Cohen, D.J.), pp. 334–56.

Tooby, J. and Cosmides, L. 1992. The psychological foundations of culture. In The Adapted Mind (ed. Barkow, J.H., Cosmides, L., and Tooby, J.). Oxford University Press.

Pinker, S. 1994. The Language Instinct. HarperCollins.

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Finlay, B.L., Darlington, R.B., and Nicastro, N. 2001. Developmental structure in brain evolution. Behavioral and Brain Sciences 24:263–308.

Barton, R.A. and Harvey, P.H. 2000. Mosaic evolution of brain structure in mammals. Nature 405:1055–8.

Fodor, J. 2001. The Mind Doesn’t Work That Way. MIT Press.

Pinker, S. 1997. How the Mind Works. Norton.

Lee, D. 1987. Introduction to Plato. The Republic. Penguin.

Neville-Sington, P. and Sington, D. 1993. Paradise Dreamed: How Utopian Thinkers Have Changed the World. Bloomsbury.


Conversation with the author, Montreal, 2002.

Galton, F. 1869. Hereditary Genius.

Candolle, A. de. 1872. Histoire des sciences et des savants depuis deux siècles.

Galton, F. 1874. English Men of Science: Their Nature and Nurture.

The Tempest, Act 4, scene 1.

The text of Mulcaster’s “Positions” can be found at

A Midsummer Night’s Dream, Act 3, scene 2.

Galton, F. 1875. The history of twins, as a criterion of the relative powers of nature and nurture. Fraser’s Magazine 12:566–76.

Gilham, N. 2001. A Life of Sir Francis Galton: From African Exploration to the Birth of Eugenics. Oxford University Press.

Ridley, M. 1999. Genome. 4th Estate.

Lifton, R.J. 1986. The Nazi Doctors. Basic Books.

Wright, W. 1999. Born That Way. Routledge.

For an excellent summary of the ins and outs of twins, see Segal, N. 1999. Entwined Lives. Dutton. Incidentally, it is increasingly unfashionable to use the terms “identical” and “fraternal”; researchers prefer the more precise “monozygotic” and “dizygotic.” But this is a popular book, so I stick to the popular terms.

For a general review of behavior genetics, see: Plomin, R., DeFries, J.C., Craig, I.W., and McGuffin, P. 2002. Behavioral Genetics in the Postgenomic Era. American Psychological Association.

Wright, W. 1999. Born That Way. Routledge.

Farber, S.L. 1981. Identical Twins Reared Apart: A Reanalysis. Basic Books.

There is evidence that fraternal twins are actually more similar genetically than siblings, because although they come from different sperm, they often come from the same maternal oocyte, two of whose pronuclei develop into eggs. This, however, only makes it more remarkable that they prove so different from each other in personality compared with identical twins.

Bouchard, T.J., McGue, M., Lykken, D., and Tellegen, A. 1999. Intrinsic and extrinsic religiousness: Genetics and environmental influences and personality correlates. Twin Research 2:88–98; Kirk, K.M., Eaves, L.J., and Martin, N. 1999. Self-transcendence as a measure of spirituality in a sample of older Australian twins. Twin Research 2:81–7. McCort, K., Bouchard, T. J., Lykken, D. T., Tellegen, A., and Keyes, M. 1999. Authoritarianism Revisited: Genetic and Environmental Influences Examined in Twins Reared Apart and Together. Personality and Individual Differences, 27:985–1014.

Nelkin, D. and Lindee, M.S. 1996. The DNA Mystique. W.H. Freeman.

Pioneer Fund Website.

Quoted in Wright, W. 1999. Born That Way. Routledge.

Pinker, S. 2002. The Blank Slate. Penguin.

Eley, T.C., Lichtenstein, P., and Stevenson, J. 1999. Sex differences in the etiology of aggressive and nonaggressive antisocial behavior: Results from two twin studies. Child Development 70:155–68.

Mischel, W. 1981. Introduction to Personality. Holt, Rinehart and Winston.

Thomas Bouchard, interview.

Clark, W.R. and Grunstein, M. 2000. Are We Hard-Wired? The Role of Genes in Human Behavior. Oxford University Press.

Bouchard, T.J. Jr. 1999. Genes, environment and personality, pp. 98–103 in The Nature–Nurture Debate (ed. Ceci, S.J. and Williams, W.M.). Blackwell.

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Randolph Nesse, e-mail. See also Srijan, S., Nesse, R.M., Stoltenberg, S.F., Li, S., Gleiberman, L., Chakravarti, A., Weder, A.B., and Burmeister, M. 2002. A BDNF coding variant is associated with the NEO personality inventory domain neuroticism, a risk factor for depression. Neuropsychopharmacology (in press). Originally published Aug. 27, 2002 at

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Zach Mainen, interview.

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Thompson, P.M., Cannon, T.D., Narr, K.L., van Erp, T., Poutanen, V.-P., Huttunen, M., Lohnqvist, J., Standertskjold-Nordenstam, C.-G., Kaprio, J., Khaledy, M., Dail, R., Zoumalan, C.I., and Toga, A.W. 2001. Genetic influences on brain structure. Nature Neuroscience 4:1253–8; Posthuma, D., De Geus, E.J., Baare, W.F., Hulshoff Pol, H.E., Kahn, R.S., and Boomsma, D.I. 2002. The association between brain volume and intelligence is of genetic origin. Nature Neuroscience 5:83–4.

Turkheimer, E., Haley, A., D’Onofrio, B., Waldron, M., Emery, R.E., and Gottesman, I. 2001. Socioeconomic status modifies heritability of intelligence in impoverished children. Paper at the annual meeting of the Behavior Genetics Association, Cambridge, July 2001.

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Eley, T., interview.

Dickens, W.T. and Flynn, J.R. 2001. Heritability estimates versus large environmental effects: The IQ paradox resolved. Psychological Review 108:346–69.

Williams, A.G., Rayson, M.P., Jubb, M., World, M., Woods, D.R., Hayward, M., Martin J., Humphries, S.E., and Montgomery, H.E. 2000. The ACE gene and muscle performance. Nature 403:614.

Ridley, M. 1993. The Red Queen. Penguin.

Radcliffe-Richards, J. 2000. Human Nature after Darwin. Routledge.

Flynn, J.R. (unpublished). The history of the American mind in the 20th century: A scenario to explain IQ gains over time and a case for the irrelevance of g.

For those curious about Galton’s unpublished novel, a fuller précis is given in Nicholas Gilham’s biography of Galton, cited above.


James, W. 1890. Principles of Psychology.

Quoted in Shorter, E. 1997. A History of Psychiatry. John Wiley and Sons.

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James, O. 2002. They F*** You Up: How to Survive Family Life. Bloomsbury.

The psychiatrist and writer Randolph Nesse calls this the central error of psychiatric research.

Cited in Torrey, E.F. 1988. Surviving Schizophrenia: A Family Manual. Harper and Row.

Shorter, E. 1997. A History of Psychiatry. John Wiley and Sons.

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Mirnics, K., Middleton, F.A., Lewis, D.A., and Levitt, P. 2001. Analysis of complex brain disorders with gene expression microarrays: Schizophrenia as a disease of the synapse. Trends in Neurosciences 24:479–86.

Tsuang, M., Stone, W.S., and Faraone, S.V. 2001. Genes, environment, and schizophrenia. British Journal of Psychiatry 178 (supplement 40): S18–S24.

Mednick. S.A., Machon, R.A., Huttunen, M.O., Bonett, D. 1988. Adult schizophrenia following prenatal exposure to an influenza epidemic. Archives of General Psychiatry 45:189–92; Munk-Jorgensen, P. and Ewald, H. 2001. Epidemiology in neurobiological research: Exemplified by the influenzaschizophrenia theory. British Journal of Psychiatry 178 (supplement 40):s30–S32.

Davis, J.O., Phelps, J.A., and Bracha, H.S. 1999. Prenatal development of monozygotic twins and concordance for schizophrenia. In The Nature–Nuture Debate (ed. Ceci, S.J. and Williams, W.W.). Blackwell.

Tsuang, M., Stone, W.S., and Faraone, S.V. 2001. Genes, environment, and schizophrenia. British Journal of Psychiatry 178 (supplement 40): S18–S24.

Deb-Rinker, P., Klempan, T.A., O’Reilly, R.L., Torrey, E.F., and Singh, S.M. 1999. Molecular characterization of a MSRV-like sequence identified by RDA from monozygotic twin pairs discordant for schizophrenia. Genomics 61:133–44.

Karlsson, H., Bachmann, S., Schroder, J., McArthur, J., Torrey, E.F., and Yolken, R.H. 2001. Retroviral RNA identified in the cebrebrospinal fluids and brains of individuals with schizophrenia. Proceedings of the National Academy of Sciences 98:4634–9.

Impagatiello, F., Guidotti, A.R., Pesold, C. et al. 1998. A decrease of reelin expression as a putative vulnerability factor in schizophrenia. Proceedings of the National Academy of Sciences 95:15718–23.

Fatemi, S.H., Emamian, E.S., Kist, D., Sidwell, R.W., Nakajima, K., Akhter, P., Shier, A., Sheikh, S., and Bailey, K. 1999. Defective corticogenesis and reduction in reelin immunoreactivity in cortex and hippocampus of prenatally infected neonatal mice. Molecular Psychiatry 4:145–54.

Fatemi, S.H. 2001. Reelin mutations in mouse and man: From reeler mouse to schizophrenia, mood disorders, autism, and lissencephaly. Molecular Psychiatry 6:129–33.

Hong, S.E., Shugart, Y.Y., Huang, D.T., Shahwan, S.A., Grant, P.E., Hourihane, J.O., Martin, N.D., and Walsh, C.A. 2000. Autosomal recessive lissencephaly with cerebellar hypoplasia is associated with human RELN mutations. Nature Genetics 26:93–6.

Cannon, M., Caspi, A., Moffitt, T.E., Harrington, H., Taylor, A., Murray, R.M., and Poulton, R. 2002. Evidence for early-childhood, pan-developmental impairment specific to schizophreniform disorder: Results from a longitudinal birth cohort. Archives of General Psychiatry 59:449–56.

Weinberger, D.R. 1987. Implications of normal brain development for the pathogenesis of schizophrenia. Archives of General Psychiatry 44:660–9; Weinberger, D.R. 1995. From neuropathology to neurodevelopment. Lancet 26:552–7.

Mirnics, K., Middleton, F.A., Lewis, D.A., and Levitt, P. 2001. Analysis of complex brain disorders with gene expression microarrays: Schizophrenia as a disease of the synapse. Trends in Neurosciences 24:479–86.

Horrobin, D. 2001. The Madness of Adam and Eve. Bantam.

Peet, M., Glen, I., and Horrobin, D. 1999. Phospholipid Spectrum Disorder in Psychiatry. Marius Press.

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Quoted in Horrobin, D. 2001. The Madness of Adam and Eve. Bantam.

Stevens, A. and Price, J. 2000. Prophets, Cults, and Madness. Duckworth, London.

Simonton, D.K. 2002. The Origins of Genius. Oxford University Press.

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For example, Oyama, S. 2000. Evolution’s Eye. Duke University Press.

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