phenotype

The Extended Phenotype: The Long Reach of the Gene

by Richard Dawkins  · 1 Jan 1982  · 506pp  · 152,049 words

and the Journal of Aesthetics and Art Criticism. His most recent book is Brainchildren: Essays on Designing Minds (MIT Press and Penguin, 1998). The Extended Phenotype The Long Reach of the Gene Richard Dawkins With a new afterword by Daniel Dennett Oxford University Press, Great Clarendon Street, Oxford OX2 6DP Oxford

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Modifiers 9 Selfish DNA, Jumping Genes, and a Lamarckian Scare 10 An Agony in Five Fits 11 The Genetical Evolution of Animal Artefacts 12 Host Phenotypes of Parasite Genes 13 Action at a Distance 14 Rediscovering the Organism Afterword by Daniel Dennett References Further Reading Glossary Author Index Subject Index Acknowledgements

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tend to have low heritability (Falconer 1960); ‘…evolution by natural selection destroys the genetic variance on which it feeds’ (Lewontin 1979b). Functional hypotheses frequently concern phenotypic traits, like possession of eyes, which are all but universal in the population, and therefore without contemporary genetic variation. When we speculate about, or make

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increase came about through evolution, probably propelled by natural selection. (4) Whether propelled by selection or not, at least part of the evolutionary change in phenotype reflected an underlying genetic change: allele replacement took place and consequently mean mental ability increased over generations. (5) By definition therefore, at least in the

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change. In this sense, the substitution of a neutral allele would not constitute evolution …’ If a whole-organism biologist sees a genetically determined difference among phenotypes, he already knows he cannot be dealing with neutrality in the sense of the modern controversy among biochemical geneticists. He might, nevertheless, be dealing with

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a neutral character in the sense of an earlier controversy (Fisher & Ford 1950; Wright 1951). A genetic difference could show itself at the phenotypic level, yet still be selectively neutral. But mathematical calculations such as those of Fisher (1930b) and Haldane (1932a) show how unreliable human subjective judgement can

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manifested in changed selection pressures. We have now added the more subtle point that changes in the environment may change the very nature of the phenotypic character we set out to explain. Historical constraints The jet engine superseded the propeller engine because, for most purposes, it was superior. The designers

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generation. In this race of Rana esculenta bodies, therefore, ridibunda genes are germ-line replicators, lessonae genes dead-end ones. Dead-end replicators can exert phenotypic effects. They can even be naturally selected. But the consequences of that natural selection are irrelevant to evolution (see Chapter 5). To make the next

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being unable to cross over. The ‘experience’ of any part of the ‘inversion supergene’ therefore repeatedly includes the other parts of the supergene and their phenotypic consequences. A habitat selection gene anywhere in such a supergene, say a gene that makes individuals choose dry microclimates, would then provide a consistent habitat

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survival and reproduction. Segregation distorters are therefore, on the whole, likely to be outlaws: we expect that selection will favour genes at other loci whose phenotypic effect is to reduce the segregation distortion. This brings us to the topic of modifiers. Modifiers The classic proving ground for the theory of modifier

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they are passively carried. The passivity of their genotypes may be an immediate consequence of the lack of cytoplasm in spermatozoa: a gene cannot achieve phenotypic expression except via cytoplasm. This is a proximal explanation. But it is at least worth toying with reversing the proposition to obtain an ultimate functional

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explanation: sperms are made small, as an adaptation to prevent the phenotypic expression of the haploid genotype. On this hypothesis we are proposing an arms race between (haploid-expressed) genes for increased competitive ability among spermatozoa on

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justified in expecting to see genes assisting molecularly different alleles at their own locus within a species gene-pool, provided they had the same phenotypic effects. A phenotypically neutral mutation at a locus changes molecular identity but does nothing to weaken any selection there may be in favour of mutual assistance. Green

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may be transcribed into RNA, but then ‘spliced out’ before the RNA is translated into amino acid sequences. Either way, it is never expressed phenotypically, if by phenotypic expression we mean expression via the orthodox route of controlling protein synthesis (Doolittle & Sapienza 1980). This does not mean, however, that the so-called

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properties be? Paradoxically, we are most familiar with the more indirect, elaborate, and roundabout methods by which DNA molecules secure their future. These are their phenotypic effects on bodies, achieved by the proximal route of controlling protein synthesis, and hence by the more distal routes of controlling embryonic development of morphology

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to this kind of selection, although here the intragenomic selection pressures will probably be swamped by more powerful pressures, positive and negative, resulting from conventional phenotypic effects. Conventional selection results in changes in the frequency of replicators relative to their alleles at defined loci on the chromosomes of populations. Intragenomic selection

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, forward direction. This is why bodily adaptations can come about by selection. Genes are allowed to exert their normal effects on development. Their developmental consequences—phenotypic effects—feed back on those genes’ chances of surviving, and as a result gene frequencies change in succeeding generations in adaptive directions. Selective theories of

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adaptation, but not instructive theories, can cope with the fact that the relationship between a gene and its phenotypic effect is not an intrinsic property of the gene, but a property of the forward developmental consequences of the gene when interacting with the consequences

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-nuclear inheritance. It is becoming increasingly clear that non-nuclear genes, either in organelles such as mitochondria or loose in cytoplasm, exert noticeable effects on phenotypes (Grun 1976). I had intended to include a section called The Selfish Plasmagene, discussing the expected consequences of selection acting on cytoplasmic replicators, and the

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than an ‘external’ behavioural loop. Indeed, a geneticist in the pure sense of the word need not care about the detailed pathway from gene to phenotypic effect. Strictly speaking, a geneticist who concerns himself with these interesting matters is temporarily wearing the hat of an embryologist. The pure geneticist is concerned

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elaborate and spectacular bowers (Gilliard 1963). It is as though some species have shifted part of the burden of adaptation from bodily phenotype to extended phenotype. So far the phenotypic effects we have been considering have extended only a few yards away from the initiating genes, but in principle there is no reason

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a relative complication. Fundamentally what is going on is that genes, compared with their alleles, exert quantitative, mutually interacting, mutually modifying, effects on a shared phenotype, the mound. They do this proximally by controlling the chemistry of cells in worker bodies, and hence worker behaviour. The principle is the same, whether

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internal structural feature—for termite mounds are like bodies with a complex ‘organ’ structure. How could we do a genetic study of such group-manufactured phenotypes? We need not hope to find normal Mendelian inheritance with simple dominance. An obvious complication, as already mentioned, is that the genotypes of the individuals

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to explain analogously the principles of how a science of ‘extended genetics’ might work. Using these assumptions, then, we can write down the expected extended phenotypes, considering mud colour only, resulting from crosses between the various possible founding pair genotypes. For instance, all colonies founded by a heterozygous king and a

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a working hypothesis we might expect that disputes resulting from genetic heterogeneity in termites would be resolved by similar rules. In this way the extended phenotype could take up a discrete and regular shape, despite being built by genetically heterogeneous workers. The analysis of artefacts given in this chapter seems,

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symbiotic cellulose-digesting microorganisms in the gut—flagellates or bacteria. The symbionts are obligately dependent on the termites, and the termites on them. The proximal phenotypic power of the symbiont genes is exerted via protein synthesis in symbiont cytoplasm. But just as termite genes reach out beyond the cells that enclose

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closely related individuals, members of a kin-group. The logic of the argument now seems to compel us to contemplate the possibility of an extended phenotype’s being jointly manipulated, not necessarily cooperatively, by genes from distantly related individuals, individuals of different species, even different kingdoms. This is the direction

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genes ‘for’ changes in host physiology. The symptoms of parasitic castration, sex-change, increased size, or whatever they may be, are properly regarded as extended phenotypic manifestations of parasite genes. The alternative to Baudoin’s interpretation is that changes in host physiology and behaviour are not parasite adaptations, but simply dull

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To recapitulate that story in slightly different terms, a student of snail genetics and a student of fluke genetics might each look at the same phenotypic variation, variation in snail shell thickness. The snail geneticist would partition the variance between a genetic and an environmental component, by correlating the thickness of

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the cytoplasm surrounding the nucleus in which the gene sits. Messenger RNA streams through the nuclear membrane and mediates genetic control over cytoplasmic biochemistry. The phenotypic expression of a gene is then, in the first place, its influence on cytoplasmic biochemistry. In its turn, this influences the form and structure

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‘We have here simple Mendelian inheritance the expression of which is constantly delayed one generation.’ The phenomenon perhaps arises when the embryological event determining the phenotypic trait occurs so early in development as to be influenced by maternal messenger RNA from the egg cytoplasm, before the zygote has begun to manufacture

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it by defeating all available alternative alleles. The weapons with which they won, and the weapons with which their rivals lost, are their respective phenotypic consequences. These phenotypic consequences are conventionally thought of as being restricted to a small field around the replicator itself, its boundaries being defined by the body wall

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quote key passages from the summary of Mayr’s chapter (pp. 295–296), showing how they may be adapted to the world of the extended phenotype. The phenotype is the product of the harmonious interaction of all genes. The genotype is a ‘physiological team’ in which a gene can make a maximum

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substances which ultimately benefit other cells that do contain germ-line copies of the leaf genes, the genes which gave the leaves their characteristically leafy phenotypes. But we cannot accept the conclusion of the previous paragraph, that inter-leaf vehicle selection, and inter-organ selection generally, could go on if

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is a phenomenon which has emerged as a result of natural selection on primitively independent selfish replicators. It has paid replicators to behave gregariously. The phenotypic power by which they ensure their survival is in principle extended and unbounded. In practice the organism has arisen as a partially bounded local concentration

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about the conditions under which content can be ascribed to mechanisms. In philosophical jargon, pure extensionality reigns in genetics, and this makes any labelling of phenotypic traits ‘a matter of arbitrary convenience’ but not, for that reason, unmotivated by our interest in drawing attention to the most telling facts about the

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34, 247–278. Cavalier-Smith, T. (1980). How selfish is DNA? Nature 285, 617–618. Cavalli-Sforza, L. & Feldman, M. (1973). Cultural versus biological inheritance: phenotypic transmission from parents to children. Human Genetics 25, 618–637. Cavalli-Sforza, L. & Feldman, M. (1981). Cultural Transmission and Evolution. Princeton, N.J.: Princeton University

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in Ethology (eds P. P. G. Bateson & R. A. Hinde), pp. 7–54. Cambridge: Cambridge University Press. Dawkins, R. (1978a). Replicator selection and the extended phenotype. Zeitschrift für Tierpsychologie 47, 61–76. Dawkins, R. (1978b). What is the optimon? University of Washington, Seattle, Jessie & John Danz Lecture, unpublished. Dawkins, R. (1979a

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. L. (1980). The Bruce effect: an evaluation of male/female advantages. American Naturalist 114, 932–938. Seger, J. A. (1980). Models for the evolution of phenotypic responses to genotypic correlations that arise in finite populations. PhD thesis, Harvard University, Cambridge, Mass. Shaw, G. B. (1921). Back to Methuselah. Reprinted 1977. Harmondsworth

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preformationism, implies that embryonic development is fundamentally, and in principle, irreversible (see central dogma). epistasis A class of interactions between pairs of genes in their phenotypic effects. Technically the interactions are non-additive which means, roughly, that the combined effect of the two genes is not the same as the sum

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. See also segregation distorter. meme A unit of cultural inheritance, hypothesized as analogous to the particulate gene, and as naturally selected by virtue of its ‘phenotypic’ consequences on its own survival and replication in the cultural environment. Mendelian inheritance Non-blending inheritance by means of pairs of discrete hereditary factors (now

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is often taken to culminate in the production of the adult, but strictly it includes later stages such as senescence. The doctrine of the extended phenotype would lead us to generalize ‘ontogeny’ to include the ‘development’ of extracorporeal adaptations, for example artefacts like beaver dams. optimon The unit of natural

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consider it diploid for most purposes. Chapter 11 suggests that although individual termites are diploid, the whole termite nest may be regarded as the extended phenotypic product of a tetraploid genotype. vehicle Used in this book for any relatively discrete entity, such as an individual organism, which houses replicators (q.

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, 11, 186, 195 cellular ecology, 222–223 cellular parasites, 226 central dogma, 97 of embryology, 173–176 of molecular genetics, 168 central theorem of extended phenotype, 233, 248 of sociobiology, 5, 55, 58, 233 centriole, 160 Cepaea nemoralis, 31 characteristic length, 89 chemical gradients, 203 Cheshire Cat, 223 chess, computer,

Beyond Inheritance: Our Ever-Mutating Cells and a New Understanding of Health

by Roxanne Khamsi;  · 21 Apr 2026  · 335pp  · 91,958 words

/​12/​29/​health/​research/​29cancer.html. GO TO NOTE REFERENCE IN TEXT coaxed to become tame: V. M. Weaver et al., “Reversion of the Malignant Phenotype of Human Breast Cells in Three-Dimensional Culture and In Vivo by Integrin Blocking Antibodies,” Journal of Cell Biology 137, no. 1 (1997): 231–45

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REFERENCE IN TEXT all of them had PNH clones: David J. Araten et al., “Clonal Populations of Hematopoietic Cells with Paroxysmal Nocturnal Hemoglobinuria Genotype and Phenotype Are Present in Normal Individuals,” Proceedings of the National Academy of Sciences of the United States of America 96, no. 9 (1999): 5209–14, doi

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/​10.1038/​347434a0. GO TO NOTE REFERENCE IN TEXT found ten specific DNA changes: Amanda Padovan et al., “Differences in Gene Expression Within a Striking Phenotypic Mosaic Eucalyptus Tree That Varies in Susceptibility to Herbivory,” BMC Plant Biology 13, no. 1 (2013): 29, doi.org/​10.1186/​1471-2229-13-29

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. GO TO NOTE REFERENCE IN TEXT cells with restored dystrophin: Eric P. Hoffman et al., “Somatic Reversion/Suppression of the Mouse mdx Phenotype In Vivo,” Journal of the Neurological Sciences 99, no. 1 (1990): 9–25, doi.org/​10.1016/​0022-510x(90)90195-s. GO TO NOTE

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IN TEXT The reanalysis revealed: Laura K. Jackson et al., “Helicobacter pylori Diversification During Chronic Infection Within a Single Host Generates Sub-Populations with Distinct Phenotypes,” PLOS Pathogens 16, no. 12 (2020): e1008686, doi.org/​10.1371/​journal.ppat.1008686. GO TO NOTE REFERENCE IN TEXT were more adept at colonizing

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.1046/​j.1365-2958.1999.01140.x. GO TO NOTE REFERENCE IN TEXT accelerate their evolution by a hundredfold: Ahmad Shafiee et al., “Recombination and Phenotype Evolution Dynamics of Helicobacter pylori in Colonized Hosts,” International Journal of Systematic and Evolutionary Microbiology 66, no. 7 (2016): 2471–77, doi.org/​10.1099

Human Diversity: The Biology of Gender, Race, and Class

by Charles Murray  · 28 Jan 2020  · 741pp  · 199,502 words

into the story of sex differences in cognitive repertoires eventually, but as I write they represent important findings with undetermined or uncertain effects on the phenotype. The Activational Effects of Sex Hormones Almost everyone has heard about sex hormones. Both sexes have all of the major sex hormones to some

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varying with their level in the bloodstream, which changes over time. The technical term for these effects is activational. Hormones have many such effects on phenotypic differences between males and females, and the hormones are not limited to estrogen and testosterone. For example: The female advantage in social cognition. A

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autism,48 the empathy quotient,49 systemizing quotient,50 social relationships,51 and interest in children.52 The pattern of results linking testosterone to the phenotype has been striking but not dispositive. A 2015 review of the literature on early androgen exposure and sex development (first author was Melissa Hines)

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producing more robust evidence. DISSENTING VOICES The best-known and most detailed critiques of the organizational role of testosterone in particular and biological explanations of phenotypic sex differences in general are Rebecca Jordan-Young’s Brain Storm (2010), Cordelia Fine’s pair of books, Delusions of Gender (2010) and Testosterone

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is being conducted within a consensus among neuroscientists that the male brain is more lateralized than the female brain. The differences are consistent with observed phenotypic sex differences in visuospatial and verbal skills. Sex Differences in Emotional Cognition and Memory I promised that I would give you a glimpse of

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progress in understanding sex differences in emotional response because an extensive technical literature has been accumulating and because of the intriguing links between the female phenotypic advantage in certain kinds of memory and the greater female vulnerability to depression.[109] The story that is emerging has not reached the level

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of the story. Recapitulation The takeaways from this chapter’s complicated discussion can be summarized quickly: Circulating sex hormones produce easily observable differences in the phenotype. Those hormones have specific, documented effects that match up with some of the differences in personality and neurocognitive functioning discussed in chapters 2 and 3

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of the male brain has been documented by a variety of evidence about sex differences in structural connectivity and functional connectivity. These findings bear on phenotypic sex differences in visuospatial and verbal skills. Differences in the functioning of the amygdala, hypothalamus, and other regions of the limbic system appear to

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have links with phenotypic sex differences in memory and vulnerability to depression. These topics barely scratch the surface. For example, I described sex differences in memory as they

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are related to the amygdala. Researchers are now integrating the findings on the phenotypic sex differences in memory, spatial abilities, and perceptual processing (the temporal order in which a scene and its individual features are recognized) into an

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is to get past the first hurdle in thinking about race differences: to lay out the evidence that it is evolutionarily reasonable to expect that phenotypic differences among races in cognitive repertoires could be at least partly genetic and that expanding knowledge about genetic variants supports that expectation. I also

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of molecular evolution,” introduced in the late 1960s and given full expression by population geneticist Motoo Kimura in 1983.30 The neutral theory acknowledges that phenotypic evolution is driven by Darwinian natural selection. But the theory posits that the vast majority of differences at the molecular level are neutral, meaning

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years that led to anatomically modern humans, a great deal of genetic variation has arisen that confers no particular advantage or disadvantage. Perhaps SNPs have phenotypic effects, but these effects are too small to have an appreciable impact on reproductive fitness. Perhaps a mutation spread to some percentage of the

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may not go to fixation after an environmental change—the spread of any single favorable allele for a polygenic trait slows as the organism’s phenotype becomes satisfactorily adapted to the changed environment—but its frequency within the population increases. These alterations in standing variation are known as soft sweeps,

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on hundreds of the relevant sites and thereby produces a cumulatively large effect. Such changes in standing variation can reliably produce dramatic effects in the phenotype through breeding. Humans have known this for millennia, even though they didn’t know anything about alleles. Darwin begins On the Origin of Species

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blond hair, and blue eyes. Their conclusion: “Our results suggest that selection on complex traits has been an important force in shaping both genotypic and phenotypic variation within historical times.”51 Also in 2016, Daniel Schrider and Andrew Kern published a sophisticated new machine-learning technique called Soft/Hard Inference through

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indefinitely was always implausible. It is now out of the question. Differences in Allele Frequencies Within and Across Continental Population When SNPs cause differences in phenotypic traits, evidence for that role surfaces first in differences in target allele frequencies. The target allele is usually defined as one that is associated

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Continental population differences in target allele frequencies associated with personality, abilities, and social behavior are common. I am not presenting proof that those differences cause phenotypic differences, but showing you how different the situation actually facing geneticists is from the impression you may have when you hear that “race is a

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later), but the image fostered by “race is a social construct” does not apply. The raw material for investigating genetic sources of population differences in phenotypic traits consists of differences in target allele frequencies. For subpopulations within continents, the raw material is meager. For continental populations, the raw material is

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they will in fact be investigated. It is implausible to expect that none of the imbalances will yield evidence of significant genetic differences related to phenotypic differences across continental populations. The results will often be complex. The same SNPs that affect the trait under investigation will typically be correlated with many

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pressures. Those environmental pressures have typically been confined to populations in specific geographic areas. Most recently, the task of assembling the genetic story for specific phenotypic traits has begun. It is still in its early stages, but progress is accelerating nonlinearly. Hence the nervousness that has prevented open discussion of

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of heritability is unrecognizably different. Expressed in words, heritability is a ratio calculated as the variance attributable to genes divided by total variance in the phenotype. Mathematically, the kind of heritability that I will be discussing, narrow heritability, is denoted as h2. A MINI-INTERLUDE I don’t need to

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influences that siblings do not share. In 1987, Robert Plomin, one of the leading students of the nonshared environment, suggested five unshared sources of such phenotypic differences.9 One is a catch-all nonsystematic category (e.g., accidents, illnesses, trauma). Four others are systematic: family composition (e.g., birth order,

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times the independent role of personality, the school environment, or parent-reported behavior problems.33 IQ was much more important than self-efficacy in explaining phenotypic variance, several times more important than personality, the school environment, or parent-reported behavior problems. The independent roles of the home environment, well-being,

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caused temporary changes in gene expression (as in a broken ankle), but changed the methylation patterns, thereby causing permanent genetic changes that damage the phenotype. Suppose that a subsequent positive environmental event could demethylate and thereby reactivate the genes that had been turned off by the negative event. Suppose—and

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-drinking example illustrates just how thoroughly the old jigsaw-puzzle metaphor has been blown up. The process of mapping causal chains from genetic variation to phenotypic trait is immensely more complicated than that. The Great Debate Immensely more complicated, yes. But is it impossibly complicated? Seen from another perspective, the

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by which the polygenic score affects the trait.”28 For his part, Turkheimer does not dispute the existence of the correlations between polygenic scores and phenotypic traits that Plomin describes. Yet these two schools nonetheless represent radically different understandings of where genomics and neuroscience are going to take us. The

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as of 2020 (e.g., parental SES, education, IQ), researchers have access to polygenic scores for various aspects of criminality. They analyze how the phenotypic measures interact with the polygenic measures as predictors of criminal behavior. In light of the Turkheimer school’s objections, can the researchers be sure that

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effects, and the contamination is rightly feared to be worst for people who have come from the most disadvantaged environments. Correlations between polygenic scores and phenotypes cannot be explained by backward causation, and that alone is enough to give us important leverage, despite all the complications.36 Eric Turkheimer has

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by sex and by ancestral population. Like most biological classifications, these groups have fuzzy edges. This complicates things analytically, but no more than that. Many phenotypic differences in personality, abilities, and social behavior that we observe between the sexes, among ancestral populations, and among social classes have a biological component. Growing

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good to bad, but that tend to be complementary. Neuroendocrinologist Geert de Vries has argued that sex differences in brain structure may work to prevent phenotypic differences. “Intuition tells us that sex differences in brain structure beget sex differences in brain function,” he wrote in 2005. “There is nothing wrong

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that these myriad differences must have implications at many levels of brain function. But our understanding of the specifics, and what those differences mean for phenotypic traits, is still rudimentary. Generally Greater Male Variance If you followed the furor about James Damore’s internal memo at Google that got him

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study. The meta-analysis focused on which regions were activated, with broad characterizations of the results rather than specific hypotheses about how they related to phenotypic differences. Here is the authors’ overall conclusion (omitting references embedded in the text): Here we have assembled 56 human functional imaging studies that each

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6. One of the first systematic evaluations of population stratification using polygenic scores (first author was Alicia Martin) calculated polygenic scores for eight well-studied phenotypes and concluded that polygenic scores based on a single-ancestry population have numerous problems. For example, polygenic scores based on a European sample predict that

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papers on measuring degrees of relatedness in 1921. For an accessible discussion of them, see Hill (1995). 5. For a nice illustration of correlations on phenotypic outcomes for different degrees of relatedness, see Cesarini and Visscher (2017): Fig. 1. 6. I adapted this example from Turkheimer, Pettersson, and Horn (2014):

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519. “A heritability coefficient represents the proportion of phenotypic variability that is associated with variability in genotype. As such, it is an effect size, a variance ratio, an R2 coefficient; and like any variance

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Honk, Schutter, Hermans et al. (2004). 28. Lewontin (1970). 29. Jencks (1979). 30. My conjecture was inspired by Cheverud’s conjecture that genetic correlations and phenotypic correlations are similar. Cheverud (1984); Cheverud (1988). Subsequent research (e.g., Dochtermann (2011); Sodini, Kemper, Wray et al. (2018)) indicates that the conjecture is

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the Souls of Today’s Students. New York: Simon & Schuster. Boehmer, Annemie L. M., Hennie Brüggenwirth, Cissy van Assendelft et al. 2001. “Genotype Versus Phenotype in Families with Androgen Insensitivity Syndrome.” Journal of Clinical Endocrinology and Metabolism 86 (9): 4151–60. Bolnick, Deborah A. 2008. “Individual Ancestry Inference and the

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M. 1984. “Quantitative Genetics and Developmental Constraints on Evolution by Selection.” Journal of Theoretical Biology 110 (2): 155–71. . 1988. “A Comparison of Genetic and Phenotypic Correlations.” Evolution 42 (5): 958–68. Ciarrochi, Joseph V., Amy Y. C. Chan, and Peter Caputi. 2000. “A Critical Evaluation of the Emotional Intelligence

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and a Male Brain: Morphology Versus Functionality.” Proceedings of the National Academy of Sciences 113 (14): E1971. Goldberg, Lewis R. 1993a. “The Structure of Phenotypic Personality Traits.” American Psychologist 48 (1): 26–34. Goldberg, Steven. 1974. The Inevitability of Patriarchy: Why the Biological Difference Between Men and Women Always Produces

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Interactions for Reading Achievement.” PhD dissertation, Florida State University. Sodini, Sebastian M., Kathryn E. Kemper, Naomi R. Wray et al. 2018. “Comparison of Genotypic and Phenotypic Correlations: Cheverud’s Conjecture in Humans.” Genetics 209 (3): 941–48. Sohail, Mashaal, Robert M. Maier, Andrea Ganna et al. 2018. “Signals of Polygenic

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Speiser, Phyllis W., Eric S. Knochenhauer, Didier Dewailly et al. 2000. “A Multicenter Study of Women with Nonclassical Congenital Adrenal Hyperplasia: Relationship Between Genotype and Phenotype.” Molecular Genetics and Metabolism 71: 527–34. Spengler, Marion, Juliana Gottschling, Elisabeth Hahn et al. 2018. “Does the Heritability of Cognitive Abilities Vary as a

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“Socioeconomic Status Modifies Heritability of IQ in Young Children.” Psychological Science 14 (6): 623–28. Turkheimer, Eric, Erik Pettersson, and Erin E. Horn. 2014. “A Phenotypic Null Hypothesis for the Genetics of Personality.” Annual Review of Psychology 65 (1): 515–40. Turkheimer, Eric, and Mary Waldron. 2000. “Nonshared Environment: A Theoretical

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3 (2): 159–80. Winkler, Anderson M., Peter Kochunov, John Blangero et al. 2010. “Cortical Thickness or Grey Matter Volume? The Importance of Selecting the Phenotype for Imaging Genetics Studies.” NeuroImage 53 (3): 1135–46. Witchel, Selma Feldman, and Ricardo Azziz. 2010. “Nonclassic Congenital Adrenal Hyperplasia.” International Journal of Pediatric Endocrinology

The Genetic Lottery: Why DNA Matters for Social Equality

by Kathryn Paige Harden  · 20 Sep 2021  · 375pp  · 102,166 words

narrowing of scope provides an essential qualification for all of the empirical results that I describe in the book. Genetic research on social and behavioral phenotypes, with its current focus on people of European genetic ancestry, cannot meaningfully inform our scientific understanding of social inequalities between racial and ethnic groups.

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3.1). This composite is a polygenic index. FIGURE 3.1.  Creating a polygenic index. Figure reproduced from Daniel W. Belsky and K. Paige Harden, “Phenotypic Annotation: Using Polygenic Scores to Translate Discoveries from Genome-Wide Association Studies from the Top Down,” Current Directions in Psychological Science 28, no. 1 (February

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2019): 82–90, https://doi.org/10.1177/0963721418807729. Correlations between individual SNPs and a phenotype are estimated in a “Discovery GWAS” with a large sample size. Many GWAS have samples that exceed millions of people. Then, a new person’s

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single sub-continental group, with African ancestry populations showing the greatest genetic diversity. As a consequence, the genetic variants that are most important for a phenotype in one population are not necessarily the most important in another population: a particular mutation in the CFTR gene, for instance, is responsible for over

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polygenic indices. We can’t assume that everyone who has the same race shares the same genetic ancestry. Whether we are talking about complicated social phenotypes like education or relatively uncontroversial physical ones like height—modern molecular genetic studies, like the older twin studies, have told us a whole lot

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previous chapter, outcomes like height (not to mention more complicated social outcomes like education) are influenced by many genes, which are probabilistically related to the phenotype. We typically observe probabilities by studying the frequency of certain outcomes in groups of people who lived in a particular time and place. As a

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In fact, high heritability implies that children of the same parents will diverge in their life outcomes. Heritability is about whether genetically different people show phenotypic differences, and siblings are genetically different. The Heritability of Seven Domains of Inequality We have discussed how sibling differences in identity-by-descent sharing could

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of genes on life outcomes is this: Are people who are more genetically different (in this case, fraternal twins compared to identical twins) also more phenotypically different? The more different fraternal twins are in a particular trait, like height, in comparison to identical twins, the higher the heritability of that

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identifying all of the causal genetic variants and measuring how much trait variation they explain.”16 We are, obviously, not there yet for any human phenotype, much less complicated ones like education. In the meantime, one method of obtaining a Goldilocks (not too big, not too small) estimate of heritability

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experiment to test whether the specific genes captured by a polygenic index cause differences in life outcomes. FIGURE 6.5.  Heritability estimates for four human phenotypes from three different methods. “Education” = educational attainment (years of formal schooling). “Age first birth” = women’s age at first childbirth. “BMI” = body mass index. “

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children are allowed to go to school regardless of their hair color. That change in social policy would break the causal chain between genotype and phenotype without directly manipulating anything about children’s genes or gene products. One does not have to edit embryonic DNA or give children pharmaceuticals to

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that are necessary for a neuron’s electrical charge. The centrality of the brain in terms of gene expression is also seen for every other phenotype relevant to social inequality—subjective well-being and depression, alcohol use and smoking, obesity and income. Returning again to Jencks’s example of red

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the time children were 2 years old. Other research has used polygenic indices created from the GWAS of educational attainment in order to see what phenotypes are correlated with polygenic indices, and when in development these correlations are apparent. This work has shown that the education polygenic index is correlated with

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or more generally as “non-cognitive” skills. The label “non-cognitive” is a misnomer: behavioral control and interpersonal skills are obviously brain-based, cognitively demanding phenotypes. But the “non” in “non-cognitive” serves to emphasize what these motivational, behavioral, and emotional traits are not—they are not synonymous with performance on

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what most groups have found for IQ (50% to 80%). Second, researchers have taken polygenic indices created from GWAS of educational attainment and seen what phenotypes, other than cognitive test performance, those polygenic indices are correlated with in childhood and adolescence. This work has found that “education” polygenic indices are correlated

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the fifty years since, however, we have—thankfully—learned some things. Thousands upon thousands of genetic variants matter for educational attainment and other complicated human phenotypes. These genes exert their effects via largely unknown cellular processes that are happening in neurons and other brain cells. These cellular effects are already happening

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educational outcomes, would Koellinger’s probability of getting a PhD have been different if the Berlin wall had not come down? Also yes. Heritable phenotypes are not immune from social change. Unfortunately, the mistaken idea that genetic influences are an impermeable barrier to social change is also widely endorsed not

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much more complicated—questions. First, how have social and historical contexts differed in ways that, like putting on eyeglasses, change the relationship between genotype and phenotype? Second, looking forward to the question of policy, what do we want the relationship between people’s genetics and their outcomes to look like? These

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thousands of genetic variants with tiny effects and unknown mechanisms.20 To make matters even more complicated, many of these variants are also involved in phenotypes that are valued differently by society: many of the same genetic variants associated with higher educational attainment, for instance, are also associated with higher

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inequalities are). The figure also pinpoints the outcomes of two hypothetical individuals who have different genes: genotype A (circle) versus B (triangle). How the expected phenotypic outcomes of individuals with a certain genotype vary across alternative environments is the reaction norm.36 The idea of shifts in reaction norm emphasizes a

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societies. The length and complexity of this causal chain means that there are multiple opportunities to intervene in the connection between genotype and a complex phenotype. Changing the health care system so that wages for “low-skilled” workers were not dragged down by the immense cost of employer-provided health

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or height, your parents’ genes didn’t make you taller or fatter unless you actually inherited them; the untransmitted alleles were uncorrelated with the child phenotype. For education, on the other hand, your parents’ genes are still associated with your own ultimate educational attainment—even if you didn’t inherit

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to blame for being overweight,” or “Genes influence people’s mood and emotions so they are less to blame for being depressed.” Depending on the phenotype, people respond to genetic information differently. FIGURE 10.1.  Rates of criminal justice system involvement and antisocial behavior by polygenic index created from GWAS of

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I previously told you about in chapter 2. In that study, psychologists at the University of Minnesota asked people to estimate how heritable different phenotypes are.13 The people who were most accurate were mothers with more than one child, but generally folks converged on something resembling the right answer

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in Western industrialized societies. There were, however, two exceptions. In an interesting twist, people in this study substantially over-estimated the heritability of just two phenotypes—breast cancer and sexual orientation. People who described themselves as politically liberal had particularly high estimates of the heritability of sexual orientation. Breast cancer and

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ability to respond to complex circumstances in complex and unpredictable ways and in the process build the self. In Turkheimer’s view, the individual phenotypic space that is not determined by either your genotype or the environmental circumstances defines the boundaries in which your free will gets to play. To

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agency over their outcomes,18 let us consider what we actually observe about identical-twin differences in social and economic outcomes, and in the psychological phenotypes that are rewarded (in modern industrialized capitalist societies) with social and economic success. Turkheimer notes that the e2 coefficient for IQ is only a

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outcomes—scores on intelligence tests foremost among them—automatically activate notions about human inferiority and superiority. I then want to consider alternative examples of human phenotypes—such as height, deafness, and autism—where genetic research has been largely embraced rather than rejected as dangerous. Can we look to these examples

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Socially Valued, Not Inherently Valuable The tendency to see intelligence (as measured on standardized IQ tests) and educational success, perhaps more than any other human phenotypes, in terms of a hierarchy of inferior and superior persons is not an accident. It is an idea that was deliberately crafted and disseminated. As

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between inherently valuable and socially valued might be unfamiliar as applied to our understanding of intelligence test scores, but we can look to three other phenotypes where it is more typical: height, deafness, and autism spectrum disorders. In chapter 2, I told you about the towering NBA player Shawn Bradley,

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-and-effect-20180515/. 15. The evolutionary biologist Richard Dawkins made the point that genetic causes should be defined as difference makers even for relatively simple phenotypes that are intuitively “genetic,” such as eye color. He wrote, “The ‘effect’ of any would-be cause can be given meaning only in terms

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, whether or not people graduate from college. The scientific and philosophical importance of heritability statistics is derived from the scientific and philosophical importance of the phenotype. Eric Turkheimer, “Three Laws of Behavior Genetics and What They Mean,” Current Directions in Psychological Science 9, no. 5 (October 1, 2000), 160–64,

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distribution of Y for those individuals” (p. 40). Selection experiments are an interesting twist on this requirement. The claim that genes (X) cause the phenotype (Y) means that for at least some individuals, there is a possible manipulation of some value of X that they possess. In the case of

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certain values, this will change the probability distribution of Y for those individuals’ offspring. If selection experiments demonstrate the causal power of genes for the phenotype, and heritability determines the response to selection, it is impossible to conclude that heritability is somehow irrelevant to causation. As Peter Visscher described in another

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steadily mounting evidence that genes mattered for understanding social inequality. But that’s exactly what he didn’t find! Instead, the study found that twins’ phenotypic similarity (i.e., how similar twins are for their outcomes) tracked their actual genetic relationship, not what their parents thought their zygosity was—evidence in

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Inequality: A Reassessment of the Effect of Family and Schooling in America (New York: Basic Books, 1972). 2. Complicated human behaviors are not the only phenotypes that are connected to genotypes via long causal chains. As the evolutionary biologist Richard Dawkins argued, “What on earth [is] any genetic trait … morphological, physiological

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depend on long causal chains involving complex social processes, such as peer norms and teacher effects, in order to be effective. Richard Dawkins, The Extended Phenotype: The Long Reach of the Gene, rev. ed. (Oxford and New York: Oxford University Press, 1999) 3. Paul Oppenheim and Hilary Putnam, “Unity of

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Academy of Sciences 115, no. 31 (July 31, 2018): E7275–84, https://doi.org/10.1073/pnas.1801238115; Daniel W. Belsky and K. Paige Harden, “Phenotypic Annotation: Using Polygenic Scores to Translate Discoveries from Genome-Wide Association Studies from the Top Down,” Current Directions in Psychological Science 28, no. 1 (February

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(1979): 327–47, https://doi.org/10.2307/2553675. 4. Heritability does not have clear implications for whether environmentally induced change is possible for a phenotype, but it might have implications for whether those environmentally induced changes persist across generations. Returning to Goldberger’s example of eyeglasses, one’s own vision

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Thomas D. Cook et al., Sesame Street Revisited (New York: Russell Sage Foundation, 1975). 36. Anthony J. F. Griffiths et al., “Norm of Reaction and Phenotypic Distribution,” in An Introduction to Genetic Analysis, 7th ed., ed. Anthony J. F. Griffiths et al. (New York: W. H. Freeman, 2000), http://www.

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the physical causes are genetic? Why are genetic determinants thought to be any more ineluctable, or blame-absolving, than environmental ones?” Richard Dawkins, The Extended Phenotype: The Long Reach of the Gene, rev. ed. (Oxford and New York: Oxford University Press, 1999). 15. The existence of genomic differences between monozygotic

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twins means that twin estimates of heritability might be systematically underestimated, as phenotypic differences between monozygotic twins caused by genetic differences between them would be misattributed to environmental variation. Hakon Jonsson et al., “Differences between Germline Genomes of

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upper bound of the extent to which people have agency. What the neuroscientist Kevin Mitchell calls “developmental variation,” i.e., inherent randomness in processes of phenotypic development, will also pull twins away from one another, without either one of them exerting anything we would typically recognize as agency. Kevin J. Mitchell

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as about differences, 115–117; coefficient, 116; criminal behavior, 195–196; equality versus equity and, 159–164; GWAS and, 123–125; objections to, 121–123; phenotypes and, 131–132; red-headed children and, 131–136; of seven domains of inequality, 117–121; twin studies of, 117–121; when the worst environments

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70 Padden, Carol, 222–223 Papageorge, Nicholas, 41–43, 44, 189 Parasite, 231–232 Parens, Erik, 175, 213 Pearl, Judea, 103 Pearson, Karl, 13, 19 phenotype, 57, 131–132, 155 phenylketonuria (PKU), 160–161 Pioneer Fund, 15 Plomin, Robert, 15 polygenic indices, 9–10, 33, 35, 42, 43; heritability and, 122

Epigenetics Revolution: How Modern Biology Is Rewriting Our Understanding of Genetics, Disease and Inheritance

by Nessa Carey  · 31 Aug 2011  · 357pp  · 98,854 words

such as particular illnesses. They basically allow us to explore mathematically the link between the sequences of our genes (genotype) and what we are like (phenotype), be this in terms of height, health, freckles or anything else we would like to measure. This is done by calculating how often both

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who create the most intriguing scientific problem. Why do two genetically identical individuals, who in many cases have experienced very similar environments, have such variable phenotypes? Similarly, why is it quite rare for both MZ twins in a pair to develop type 1 diabetes? What is it, in addition to the

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women) These data are consistent with a model where epigenetic changes could account for at least some of the reasons why MZ twins aren’t phenotypically identical, but there’s still a lot of supposition involved. That’s because for many purposes humans are a quite hopeless experimental system. If

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we want to be able to assess the role of epigenetics in the problem of why genetically identical individuals are phenotypically different from one another, we would like to be able to do the following: Analyse hundreds of identical individuals, not just pairs of them;

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the experimenters are most interested in. A mouse of a different colour The most useful mouse model for exploring how epigenetic changes can lead to phenotypic differences between genetically identical individuals is called the agouti mouse. Normal mice have hair which is banded in colour. The hair is black at the

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The agouti mouse has provided a quite clear-cut example of how epigenetic modification, in this case DNA methylation, can make genetically identical individuals look phenotypically different. However, there is always the fear that agouti is a special case, and maybe this is a very uncommon mechanism. This is particularly of

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development lead to non-identical patterns of gene expression. These become epigenetically set and exaggerated over the years, until eventually the genetically identical twins become phenotypically different, sometimes in the most dramatic of ways. Such a random process, caused by individually minor fluctuations in the expression of epigenetic genes during early

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to identify the environmental effects that lead to some chronic human conditions. If we study pairs of MZ twins who are discordant for a specific phenotype, for example multiple sclerosis, it may be well nigh impossible to identify an environmental cause. It may simply be that one of the pair

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to have larger arm muscles than the sons of weavers (a much less physical occupation). Lamarck interpreted this as the blacksmiths’ sons inheriting the acquired phenotype of large muscles from their fathers. Our modern interpretation is different. We recognise that a man whose genes tended to endow him with the ability

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inheritance. If we wanted to summarise a century and a half of evolutionary theory in one paragraph we might say: Random variation in genes creates phenotypic variation in individuals. Some individuals will survive better than others in a particular environment, and these individuals are likely to have more offspring. These offspring

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competitors in a particular environment. This is where the Lamarckian model of acquired characteristics really falls over, relative to Darwinian models. An acquired change in phenotype would somehow have to ‘feed-back’ onto the DNA script and change it really dramatically, so that the acquired characteristic could be transmitted in the

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This scandal tainted an already controversial field1. One of the statements in our potted history of evolutionary theory was the following, ‘An acquired change in phenotype would somehow have to ‘feed-back’ onto the DNA script and change it really dramatically so that the acquired characteristic could be transmitted in the

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risk of dying as a consequence of diabetic illnesses4. Just like Camilla in the Dutch Hunger Winter example, the sons and grandsons had an altered phenotype (a change in the risk of death through cardiovascular disease or diabetes) in response to an environmental challenge they themselves had never experienced. These data

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was important, as the more data points we have in an experiment, the more we can rely on the findings. Statistical tests showed that the phenotypic differences between the genetically identical groups were highly significant. In other words, it was very unlikely that the effects occurred by chance5. The results from

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epigenetic inheritance has taken place. In other words, an epigenetic modification (probably DNA methylation) was transferred along with the genetic code. This transfer of the phenotype from one generation to the next wasn’t perfect – not all the offspring looked exactly the same as their mother. This implies that the DNA

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methylation that controls the expression of the agouti phenotype wasn’t entirely stable down the generations. This is quite analogous to the effects we see in suspected cases of human transgenerational inheritance, such as

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the different patterns of colour in his offspring. But there are other examples of epigenetic inheritance transmitted from both males and females. The kinked tail phenotype in mice, which is caused by variable methylation of a retrotransposon in the AxinFu (Axin fused) gene, can be transmitted by either the mother

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at the AxinFu gene from either parent to offspring. These model systems have been really useful in demonstrating that transgenerational inheritance of a non-genetic phenotype does actually occur, and that this takes place via epigenetic modifications. This is truly revolutionary. It confirms that for some very specific situations Lamarckian

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inheritance is taking place, and we have a handle on the molecular mechanism behind it. But the agouti and kinked tail phenotypes in mice both rely on the presence of specific retrotransposons in the genome. Are these special cases, or is there a more general effect

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examples of transgenerational inheritance of non-genetic features are the agouti mouse and the AxinFu mouse, which we met in the previous chapter. The phenotypes in both these models are a consequence of the methylation levels of an IAP retrotransposon upstream of a gene. The DNA methylation levels in the

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parent get passed on to the offspring, and so does the phenotype caused by the expression levels of the retrotransposon9. We met other examples of transgenerational inheritance of acquired characteristics in Chapter 6, including the effects of

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Figure 8.2 Two children may each have the same deletion on chromosome 15, shown schematically by the absence of the horizontally striped box. The phenotype of the two children will be different, depending on how they inherited the abnormal chromosome. If the abnormal chromosome was inherited from their father, the

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copies of this gene, rather than just one, twice as much IGF2 protein as normal is produced and the foetus grows too much. The opposite phenotype to Beckwith-Wiedemann syndrome is a condition called Silver-Russell syndrome18,19. Children with this disorder are characterised by retarded growth before and after birth

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the correct maternal mark. When this egg is fertilised by a sperm, both copies of chromosome 15 will function like paternal chromosomes, and create a phenotype just like uniparental disomy. Research is ongoing into how all these processes are controlled. We don’t fully understand how imprints are protected from reprogramming

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tail trait that we met in earlier chapters. We know that not all transgenerational inheritance happens in the same way. In the agouti mouse the phenotype is transmitted via the mother, but not via the father. In this case, the DNA methylation on the IAP retrotransposon is removed in both

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the offspring. Male agouti mice don’t pass on either DNA methylation or repressive histone modifications on their retrotransposon, which is why transmission of the phenotype only occurs through the maternal line30. This is a slightly more indirect method of transmitting epigenetic information. Instead of direct carry-over of DNA methylation

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, an intermediate surrogate (a repressive histone modification) is used instead. This is probably why the maternal transmission of the agouti phenotype is a bit ‘fuzzy’. Not all offspring are exactly the same as the mother, because there is a bit of ‘wriggle-room’ in how

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a testis-determining pathway in the embryo. This leads to production of testosterone, the archetypal ‘male’ hormone, which then masculinises the embryo. Occasionally, individuals who phenotypically appear to be girls have the male 46, XY karyotype. In these cases the SRY gene is often inactive or deleted and consequently the foetus

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develops down the default female pathway1. Sometimes, the other scenario arises. Individuals who phenotypically appear to be boys can have the typically female karyotype of 46, XX. In these cases a tiny section of the Y chromosome containing the

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the ultimate demonstration of the power of a random epigenetic event. Two identical individuals, each with two apparently identical X chromosomes, had a completely discordant phenotype, because of a shift in the epigenetic balance of power. Sometimes, however, it is essential that individual cells express the correct amount of a protein

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of gametes is severely compromised. Leaving aside the infertility, there are two obvious conclusions we can draw from this table. The first is that the phenotypes are all relatively mild compared with, for example, trisomy of chromosome 21 (Down’s syndrome). This suggests that cells can tolerate having too many or

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having extra copies of an autosome. But the other obvious conclusion is that an abnormal number of X chromosomes does indeed have some effects on phenotype. Why should this be? After all, X inactivation ensures that no matter how many X chromosomes are present, all bar one get inactivated early

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X females compared with 47, XXX females or with the normal 46, XX female constitution. Similarly, males with the normal 46, XY karyotype should be phenotypically identical to males with the 47, XXY karyotype. In all of these cases there should be only one active X chromosome in the cells. One

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we analyse the sequence of a gene in human cells, we can find a gene of broadly similar sequence in the nematode worm. So the phenotypic differences between worms and humans aren’t caused by Homo sapiens having more, different or ‘better’ genes. Admittedly, more complicated organisms tend to splice

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compensatory mechanisms, defects may only have relatively subtle impacts. The problem this creates experimentally is that most genetic screens are good at detecting the major phenotypes caused by mutations in proteins, but may not be so useful for more subtle effects. There is a small ncRNA called BC1 which is expressed

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a number of related compounds and tested them for their effects in cell culture. The ones that inhibited DNA methylation also caused the changes in phenotype originally observed for 5-azacytidine. Compounds that didn’t inhibit DNA methylation had no effect on phenotype5. The methylation cul-de-sac Cytidine (base

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to adult stress, there is probably more than one gene involved. Both the cortisol receptor gene and the arginine vasopressin gene can contribute to this phenotype in rodents. Secondly, the studies also show us that a particular class of epigenetic modification is not in itself good or bad. It’s

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of the epigenetic changes that have been observed are relatively small. The sceptics are unconvinced that such small molecular changes could lead to such pronounced phenotypes. They argue that just because the changes are present, it doesn’t mean they’re necessarily having a functional effect. They worry that the

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For honeybees the DNA script is constant but the outcome is variable. The outcome is controlled by an early event (feeding pattern) which sets a phenotype that is maintained throughout the rest of life. This is a scenario that just shrieks epigenetics at us, and in the last few years scientists

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where Dnmt3 had been knocked down were like those of the normal royal jelly-induced queens. This is what we would expect given the similar phenotypes in the two groups. The gene expression patterns in the normal queens and the Dnmt3-knockdown queens were also very similar. The authors concluded

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by epigenetic alterations in DNA and its accompanying proteins if one or both of the following conditions are met: Two things are genetically identical, but phenotypically variable; An organism continues to be influenced by an event long after this initiating event has occurred. We always have to apply a common sense

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we forget how much we can learn just by looking thoughtfully. For example, we don’t always need sophisticated laboratory equipment to determine if two phenotypically different things are genetically identical. Here are a couple of examples with which we are all familiar. Maggots turn into flies and caterpillars turn into

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In mammals, there’s usually a clear genetic reason why males are males and females are females. A functional Y chromosome leads to the male phenotype. In lots of reptile species, including crocodiles and alligators, the two sexes are genetically identical. You can’t predict the sex of a crocodile

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of the embryo . Chromatin DNA in combination with its associated proteins, especially histone proteins. Concordance The degree to which two genetically identical individuals are identical phenotypically. CpG A cytosine nucleotide followed by a guanine nucleotide in DNA. CpG motifs can undergo methylation on the C. Discordance The degree to which two

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genetically identical individuals are non-identical phenotypically. DNA replication Copying DNA to create new DNA molecules which are identical to the original. DNMT DNA methyltransferase. An enzyme that can add methyl groups

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one brain cell that acts on another brain cell to alter its behaviour. Nucleosome Combination of eight specific histone molecules with DNA wrapped around them. Phenotype The observable characteristics or traits of an organism. Pluripotency The ability of a cell to give rise to most other cell types. Typically, pluripotent mammalian

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to give rise to all cells of the body and the placenta. Transcription Copying DNA to create RNA molecules. Transgenerational inheritance The phenomenon in which phenotypic changes in one generation are passed on to the next, without any alteration in the genetic code. Uniparental disomy A situation where both members of

The Selfish Gene

by Richard Dawkins  · 1 Jan 1976  · 365pp  · 117,713 words

a gene's-eye view of nature. It is a different way of seeing, not a different theory. In the opening pages of The Extended Phenotype, I explained this using the metaphor of the Necker cube. This is a two-dimensional pattern of ink on paper, but it is perceived as

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years: Robert Axelrod's The Evolution of Cooperation, because it seems to offer some sort of hope for our future; and my own The Extended Phenotype because for me it dominated those years and because-for what that is worth-it is probably the finest thing I shall ever write. The

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shall we resolve this paradox of the two ways of looking at life? My own attempt to do so is spelled out in The Extended Phenotype, the book that, more than anything else I have achieved in my professional life, is my pride and joy. This chapter is a brief distillation

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of a few of the themes in that book, but really I'd almost rather you stopped reading now and switched to The Extended Phenotype! On any sensible view of the matter Darwinian selection does not work on genes directly. DNA is cocooned in protein, swaddled in membranes, shielded from

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likely to develop into a successful adult, an adult likely to reproduce and pass those very same genes on to future generations. The technical word Phenotype is used for the bodily manifestation of a gene, the effect that a gene, in comparison with its alleles, has on the body, via development

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. The phenotypic effect of some particular gene might be, say, green eye colour. In practice most genes have more than one phenotypic effect, say green eye colour and curly hair. Natural selection favours some genes rather than others

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not because of the nature of the genes themselves, but because of their consequences-their phenotypic effects. Darwinians have usually chosen to discuss genes whose

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phenotypic effects benefit, or penalize, the survival and reproduction of whole bodies. They have tended not to consider benefits to the

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passed on. Here the paradox conveniently disappears because what is good for one gene is good for all. But what if a gene exerted a phenotypic effect that was good for itself but bad for the rest of the genes in the body? This is not a flight of fancy. Cases

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question. The instrument with which we shall purge our minds is the idea that I call the extended phenotype. It is to this, and what it means, that I now turn. The phenotypic effects of a gene are normally seen as all the effects that it has on the body in which

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it sits. This is the conventional definition. But we shall now see that the phenotypic effects of a gene need to be thought of as all the effects that it has on the world. It may be that a gene

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as a matter of fact. It will not be something that ought to be part of our very definition. In all this, remember that the phenotypic effects of a gene are the tools by which it levers itself into the next generation. All that I am going to add is that

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the tools may reach outside the individual body wall. What might it mean in practice to speak of a gene as having an extended phenotypic effect on the world outside the body in which it sits? Examples that spring to mind are artefacts like beaver dams, bird nests and caddis

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cellular wiring up of the nervous system. The caddis house is only a further extension of this kind of sequence. Stone hardness is an extended phenotypic effect of the caddis's genes. If it is legitimate to speak of a gene as affecting the wrinkliness of a pea or the nervous

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, isn't it? Yet the reasoning is inescapable. We are ready for the next step in the argument: genes in one organism can have extended phenotypic effects on the body of another organism. Caddis houses helped us take the previous step; snail shells will help us take this one. The shell

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shell is a fluke adaptation. If it is, it has to have come about by Darwinian selection of fluke genes. We have demonstrated that the phenotypic effects of a gene can extend, not only to inanimate objects like stones, but to 'other' living bodies too. The story of the snails and

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the weight of a normal adult. No good for propagating beetle genes, but a cornucopia for Nosema parasites. Giantism in beetle larvae is an extended phenotypic effect of protozoan genes. And here is a case history to provoke even more Freudian anxiety than the Peter Pan beetles-parasitic castration! Crabs are

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, if we accept that they are Darwinian adaptations for the benefit of the parasite, must be seen as extended phenotypic effects of parasite genes. Genes, then, reach outside their 'own' body to influence phenotypes in other bodies. To quite a large extent the interests of parasite genes and host genes may coincide

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methods that they use, there is no reason why these should be any different from the machinations-all too predictable to a selfish gene/extended phenotype theorist-of viruses. When we have a cold or a cough, we normally think of the symptoms as annoying byproducts of the virus's activities

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far discussed have lived inside their hosts. The genes, then, are physically close to their extended phenotypic effects, as close as genes ordinarily are to their conventional phenotypes. But genes can act at a distance; extended phenotypes can extend a long way. One of the longest that I can think of spans a

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and eighteenth-century coal merchants used canals. Whatever its benefits, a beaver lake is a conspicuous and characteristic feature of the landscape. It is a phenotype, no less than the beaver's teeth and tail, and it has evolved under the influence of Darwinian selection. Darwinian selection has to have genetic

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beaver genes that made good lakes for transporting trees, just as it favoured genes that made good teeth for felling them. Beaver lakes are extended phenotypic effects of beaver genes, and they can extend over several hundreds of yards. A long reach indeed! Parasites, too, don't have to live inside

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, yet we have no hesitation in labelling them as parasites. Cuckoo adaptations to manipulate the behaviour of foster-parents can be looked upon as extended phenotypic action at a distance by cuckoo genes. It is easy to empathize with foster parents duped into incubating the cuckoo's eggs. Human egg collectors

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about flukes and snails we accustomed ourselves to the idea that a parasite's genes could have phenotypic effects on the host's body, in exactly the same way as any animal's genes have phenotypic effects on its 'own' body. We showed that the very idea of an 'own' body was

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any internal drug or hormone. As in the case of internal parasites, we should now rephrase the whole matter in terms of genes and extended phenotypes. In the evolutionary arms race between cuckoos and hosts, advances on each side took the form of genetic mutations arising and being favoured by natural

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the besotted host. In exactly the same sense as we may speak of cuckoo genes having (phenotypic)effects on the colour and shape of cuckoo gapes, so we may speak of cuckoo genes having (extended phenotypic) effects on host behaviour. Parasite genes can have effects on host bodies, not just when the

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has been reinvented so often. Some examples that we'll look at have gone beyond familiar cuckooism to fulfil the wildest fantasies that The Extended Phenotype might have inspired. A bird cuckoo deposits her egg and disappears. Some ant cuckoo females make their presence felt in more dramatic fashion. I don

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is an act of special genetic madness and formidable indeed must be the drug that drives them to it. In the world of the extended phenotype, ask not how an animal's behaviour benefits its genes; ask instead whose genes it is benefiting. It is hardly surprising that ants are exploited

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species. In all cases in which natural selection has favoured genes for manipulation, it is legitimate to speak of those same genes as having (extended phenotypic) effects on the body of the manipulated organism. It doesn't matter in which body a gene physically sits. The target of its manipulation may

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favours those genes that manipulate the world to ensure their own propagation. This leads to what I have called the Central Theorem of the Extended Phenotype: An animals behaviour tends to maximize the survival of the genes 'for' that behaviour, whether or not those genes happen to be in the body

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purpose of the body of an individual lion, wolf, or deer. That this is true is now widely accepted, but why is it true? Extended phenotypes and parasites can again help us. We saw that when the genes of a parasite work together with each other, but in opposition to the

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that life, as a matter of fact, is bundled into discrete, individually purposeful vehicles like wolves and bee-hives. But the doctrine of the extended phenotype has taught us that it needn't have been so. Fundamentally, all that we have a right to expect from our theory is a battleground

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of replicators, jostling, jockeying, fighting for a future in the genetic hereafter. The weapons in the fight are phenotypic effects, initially direct chemical effects in cells but eventually feathers and fangs and even more remote effects. It undeniably happens to be the case that

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these phenotypic effects have largely become bundled up into discrete vehicles, each with its genes disciplined and ordered by the prospect of a shared bottleneck of sperms

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with a single genetic exit route? Why did genes choose to gang up and make large bodies for themselves to live in? In The Extended Phenotype I attempt to work out an answer to this difficult problem. Here I can sketch only a part of that answer-although, as might be

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other. The two are mutually enhancing, like the spiralling feelings of a woman and a man during the progress of a love affair. The Extended Phenotype is a long book and its argument cannot easily be crammed into one chapter. I have been obliged to adopt here a condensed, rather intuitive

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I have succeeded in conveying the flavour of the argument. Let me end with a brief manifesto, a summary of the entire selfish gene/extended phenotype view of life. It is a view, I maintain, that applies to living things everywhere in the universe. The fundamental unit, the prime mover of

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the replicators come first, in importance as well as in history. One way to remind ourselves is to reflect that, even today, not all the phenotypic effects of a gene are bound up in the individual body in which it sits. Certainly in principle, and also in fact, the gene reaches

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a long way away. With only a little imagination we can see the gene as sitting at the centre of a radiating web of extended phenotypic power. And an object in the world is the centre of a converging web of influences from many genes sitting in many organisms. The long

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reach of the gene knows no obvious boundaries. The whole world is criss-crossed with causal arrows joining genes to phenotypic effects, far and near. It is an additional fact, too important in practice to be called incidental but not necessary enough in theory to be

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these causal arrows have become bundled up. Replicators are no longer peppered freely through the sea; they are packaged in huge colonies-individual bodies. And phenotypic consequences, instead of being evenly distributed throughout the world, have in many cases congealed into those same bodies. But the individual body, so familiar to

Richard Dawkins: How a Scientist Changed the Way We Think

by Alan Grafen; Mark Ridley  · 1 Jan 2006  · 286pp  · 90,530 words

to fame, went on to display the range and depth of his analytical skills and literary abilities in a string of best-sellers: The Extended Phenotype (intended primarily for fellow biologists), The Blind Watchmaker, River Out of Eden, Climbing Mount Improbable, Unweaving the Rainbow, and The Ancestor’s Tale. A

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my staunchest guide. Here was a Darwinian world that was gene-centred, adaptationist; this had to be how natural selection worked. That, and The Extended Phenotype, introduced me to fundamental questions of evolutionary theory. And they taught me how, holding steadily to a gene-centred view, I could find the way

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metre centipede-like creature (Arthro-pleura). We know this from fossil footprints preserved in sandstone on the Isle of Arran. Are these footprints an extended phenotype? If so, it might appear that any grass blade bent by a passing beetle would deserve the name but consequently dilute the concept. The Extended

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or, as I should say, on the frequency of alleles linked to footprint creation. Consequently, there seems no merit in calling the footprints an extended phenotype. The Surrey puma question can now be answered: the scratch marks (even allowing for media misrepresentation) were not themselves a puma or for that matter

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a dog. They were certainly the phenotypic expression of genes for scratching behaviour. Yet, in the absence of any evidence that they affect the survival of scratch-generating genes, they do not

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vigorous thrashing movements of its body. Manipulated by the parasites, the caterpillar has become their protector, even as it dies. Notice here that the extended phenotype is the collective outcome of genes dispersed among several parasites, emphasizing the difficulty of regarding the individual as the unit of selection. For me this

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about how cooperatively-built structures, such as social insect nests, might evolve. This question is addressed with great insight by Richard Dawkins in The Extended Phenotype but, as a preliminary to describing this, it may help to know something about the village of Lavenham in Suffolk. Tourists flock to the heart

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in human-built structures it is possible for design and technology to change independent of one another. What still startles me on rereading The Extended Phenotype is how clearly Richard Dawkins already understood this in 1982 and realized that it had important implications for nest building by social insects. A mature

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in material technology compared to design. It is surprising that these important speculations, and indeed others relating to animal building behaviour, arising from The Extended Phenotype have not yet stimulated the research effort they deserve. It is a book that particularly speaks to researchers; there is valuable detail in its message

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nature of the gametes is not determined by the personal qualities of the parents or ancestors in question.’ Thus, Johannsen made a crucial distinction between phenotype (observable traits) and genotype (heritable factors). In Johannsen’s view, the mistaken notion of the inheritance of personal qualities was reinforced by the persistence of

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in all cases where new or revised conceptions are being developed... . Therefore I have proposed the terms “gene” and “genotype” and some further terms, as “phenotype” and “biotype”, to be used in the science of genetics. The “gene” is nothing but a very applicable little word, easily combined with others, and

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” or “allelomorphs” in the gametes, demonstrated by modern Mendelian researches.’ From this beginning, Johannsen’s gene has had an illustrious history (as have ‘genotype’ and ‘phenotype’, but not ‘biotype’). But ‘gene’ itself conveys little information, consisting as it does of only four letters and a single syllable when spoken. The factors

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demonstrate “genotypical” differences or accordances... . genotypes can be examined only by the qualities and reactions of the organisms in question.’ Genes are known by their phenotypic effects. Johannsen was dismissive of attempts to localize genes. ‘The question of chromosomes as the presumed “bearers of hereditary qualities” seems to be an idle

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of the unit of inheritance.) Supporters of the chromosomal theory, however, continued to define the gene operationally as that which was responsible for a heritable phenotypic difference. A. H. Sturtevant, one of the first to map genes to chromosomes, commented in 1915: ‘We can ... in no sense identify a given

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many other genes’.3 Much of twentieth-century experimental genetics was engaged in making inferences about the physical nature of genes from observations of their phenotypic effects (i.e., by observations of differences in an organism’s physical characteristics). These studies led to a definition of the gene as a

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defined chemical properties. Now, the existence of a gene is often inferred from properties of a DNA sequence without any information about the gene’s phenotypic effects and without the observation of differences among sequences. But the definition of the gene as a protein-encoding stretch of DNA is more recent

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than its definition as that which is responsible for a phenotypic difference, and it is not surprising that the modern molecular definition has not fully supplanted the older operational definition. Experimental geneticists invoke genes to

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explain observed phenotypic differences. A pink-eyed fly differs from a red-eyed fly because the former possesses a gene for pink eyes inherited from both parents whereas

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the latter has inherited at least one gene for red eyes. In a similar way, evolutionary biologists often invoke genes to explain hypothetical phenotypic differences in an attempt to understand the nature of adaptation by natural selection. An ornithologist might wish to understand why males of some species help

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The extent of the strategic gene is determined by the number of replication cycles that separate the material genes responsible for the expression of a phenotypic effect from the material genes that thereby have an increased probability of being copied. Thus, the strategic gene is not a fixed entity but can

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passed on had accumulated in many steps over the course of memetic transmission. Johanssen invented the ‘gene’ to clarify the distinction between genotype (gene) and phenotype (trait). Can a similar distinction be made for a science of memetics? There are two principal kinds of things we observe that provide evidence about

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relation seems to be reversed for memetics. Memes are observed, rather than inferred from their effects, whereas their effects are in large part hidden. The phenotype/genotype distinction works fairly well for genes, but there are many unresolved problems in its application to memes. For example, let’s suppose that there

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different, overlapping ways in which a chromosome could be divided into genes. Could the same approach work for memes? Dawkins’ principal interest was in the phenotypes of organisms rather than of genes. Just as he failed to specify precisely how to divide a chromosome into genes, he did not specify how

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18 R. L. Trivers and H. Hare, ‘Haplodiploidy and the evolution of the social insects’, Science, 191 (1976): 249-263. 19 R. Dawkins, The Extended Phenotype (Oxford: W. H. Freeman, 1982). 20 See Grafen, ‘A first formal link between the Price Equation and an optimization program’ (2002) and ‘The optimisation of

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John Maynard Smith). At the same time, the controversy began increasingly to address important issues in evolutionary theory (without losing its political underpinnings). The Extended Phenotype, Dawkins’ second book,28 was one huge attempt to catch and respond to general accusations and misperceptions when it came to such things as adaptation

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often implies a particular balance of gene frequencies. 22 Segerstråle, Defenders of the Truth (2000), Chapter 10. 23 See e.g. Richard Dawkins, The Extended Phenotype (Oxford and San Francisco: W H. Freeman, 1982), 10. 24 See e.g. Ullica Segerstråle, ‘Reductionism, “Bad Science” and Politics: A Critique of Anti-

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(adaptive) function. Dawkins, just like Hamilton and Wilson, decided to concentrate on the last of Tinbergen’s famous ‘four questions’. 28 Richard Dawkins, The Extended Phenotype: The Gene as Unit of Selection (Oxford and San Francisco: W. H. Freeman, 1982). 29 Richard Dawkins, ‘Twelve misunderstandings of kin selection’, Zeitschrift fur Tierpsychologie

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defence of selfish genes’, Philosophy, October (i98ia): 562-579. 31 Richard Dawkins, The Blind Watchmaker (New York: W. W Norton, 1987). 32 Dawkins, The Extended Phenotype (1982), 19. 33 Richard C. Lewontin, Steven Rose, and Leon Kamin, Not in Our Genes (New York: Pantheon Books, 1984). 34 Richard Dawkins, ‘Sociobiology: The

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the philosophical methods and the very considerable philosophical substance of that work. Why is a philosopher writing an Afterword for this book? Is The Extended Phenotype science or philosophy? It is both; it is science, certainly, but it is also what philosophy should be, and only intermittently is: a scrupulously

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‘Evolution: The Pleasures of Pluralism’, New York Review of Books (26 June 1997), 47-52. 10 Daniel C. Dennett, ‘Afterword’, in Richard Dawkins, The Extended Phenotype (Oxford: Oxford University Press, 1999). 11 For a detailed analysis of Skinner’s position, see Daniel C. Dennett, ‘Skinner Skinned’, in Brainstorms (Cambridge, MA: MIT

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for his fellow scientists and for informed nonspecialists: his more popular books certainly cannot be considered ‘popularization’, nor is his most technical book, The Extended Phenotype, restricted to specialists. This is an example I try to emulate. A second major theme in Dawkins’ writings on life that has important parallels in

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be made by neural circuitry programming the next iteration of the movement. In the case of the evolution of genes, the actions would be extended phenotypes, the effects would be sensed as differential mortality and fecundity, and the adjustment would be made in terms of the number of descendants resulting in

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Universe (Cambridge: Cambridge University Press, 2003). 36 Conway Morris, Life’s Solution: Inevitable Humans in a Lonely Universe (2003), 196. 37 R. Dawkins, The Extended Phenotype: The Gene as the Unit of Selection (Oxford: W. H. Freeman, 1982). 38 Dawkins and Krebs, ‘Arms races between and within species’ (1979), 490. 39

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otherwise asked. Some aspects of the environment may be stable for a very long time and yet are crucial for the expression of an adaptive phenotype. Many important writers have suggested that changes in those environmental factors can produce dramatic alterations in the characteristics of an organism. This thought lay behind

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one is available.” And that reminds me to say that Laland [one of the commentators in the Biology and Philosophy issue devoted to The Extended Phenotype] has missed the irony in my apparent espousal of Bateson’s “Great Nexus of complex causal factors interacting in development”.’8 The mischievous attribution directed

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54. 5 P. Bateson [P.G.], ‘Book Review: The Selfish Gene by Richard Dawkins’, Animal Behaviour, 26 (1978): 316-318. 6 R. Dawkins, The Extended Phenotype (Oxford: W. H. Freeman, 1982). 7 M. Mameli, ‘Nongenetic selection and nongenetic inheritance’, British Journal for the Philosophy of Science, 55 (2004): 35-71. 8

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R. Dawkins, ‘Extended phenotype—but not too extended. A reply to Laland, Turner and Jablonka’, Biology and Philosophy, 19 (2004), 377-396. What's the matter with memes? Robert

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and values which distinguish one culture from another are in people’s heads. Presumably for reasons like this, Dawkins, in his second book, The Extended Phenotype, restricted a meme to being ‘a unit of information residing in the brain’.13 But we still have a conceptual problem: if replicators are restricted

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Philosophy (http://plato.stanford.edu/entries/replication/; 2001). 11 Hull and Wilson, ‘Replication’ (2001). 12 Aunger, The Electric Meme (2002). 13 Richard Dawkins, The Extended Phenotype (Oxford: Oxford University Press, 1982). 14 Aunger, The Electric Meme (2002). 15 Dan Sperber, ‘An objection to the memetic approach to culture’, in Robert Aunger

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modeling cognition and culture: Why replicators are not necessary to cultural evolution’, Journal of Cognition and Culture, 2 (2002): 87-112. 22 Dawkins, The Extended Phenotype (1982). 23 Richard Dawkins, ‘Universal Darwinism’, in D. S. Bendall (ed.), Evolution from Molecules to Men (Cambridge: Cambridge University Press, 1982), 403-425. 24 Marshall

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. Because there are unlikely to be circumstantial cues by which a female squirrel could make this distinction, the researchers inferred that she uses ‘self-referential phenotype matching’, that is, that she compares each sister to herself with respect to some genetically complex trait or traits (probably odours) and adjusts her responses

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from dubious implicit or explicit assumptions about a pre-existing, stable repertoire of paternal responses to resemblance. In a more realistic model, the baby’s phenotypic expression of potential paternity cues, the father’s perception of those cues, and the father’s reaction to those perceptions (not to mention the responses

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been psychology and linguistics. (Physics was too damned hard.) But in 1983, at the urging of the philosopher Peter Godfrey-Smith, I read The Extended Phenotype—a truly great book—and was hooked on evolutionary biology. My copy disappeared under successive waves of marginal annotations as my career irrevocably changed course

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the “levels of selection” debate away from the individual animal as the unit of evolution to the genes, and what he has called their extended phenotypes’. Simultaneously, the award description continues, Dawkins ‘applied a Darwinian view to culture through the concept of memes as replicators of culture’. Finally, ‘Dr Dawkins’ contribution

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and make them clear even for the non-scientist. I think, for example, of his recent discussion on Radio four with Jonathan Miller on extended phenotypes. Of many examples in his writings I refer to his explanation of ‘intermediates’ and why creationists are confused in calling for evidence that, by definition

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tend to think he has ‘just’ written a popular book. It is no secret that Dawkins, despite the complexity of the arguments in The Extended Phenotype, stood accused of being a scientific lightweight among his professional colleagues for having written in an accessible style. An unexpected effect of the success of

Science in the Soul: Selected Writings of a Passionate Rationalist

by Richard Dawkins  · 15 Mar 2017  · 420pp  · 130,714 words

of the book. Editor’s introduction RICHARD DAWKINS has always defied categorization. One eminent biologist of mathematical bent reviewing The Selfish Gene and The Extended Phenotype was startled to find scientific work apparently free of logical errors and yet containing not a single line of mathematics; he could come to no

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can be said to have values in the same sense. John Maynard Smith said: ‘If there were no constraints on what is possible, the best phenotype would live for ever, would be impregnable to predators, would lay eggs at an infinite rate, and so on.’ Nicholas Humphrey continues the argument with

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, and I didn’t blame her for wearily changing the subject rather than pursuing the argument. *9 I felt it necessary to begin The Extended Phenotype by admitting that it was a work of ‘unabashed advocacy’. The fact that I needed to use a word like ‘unabashed’ speaks to my point

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is epigenetic, as it is on our planet, instructive evolution cannot work. Briefly, if acquired characters are to be inherited, embryonic processes must be reversible: phenotypic change has to be read back into the genes (or equivalent). If embryology is preformationistic – the genes are a true blueprint – then it may indeed

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a cake than a blueprint for a house, it is irreversible. There is no one-to-one mapping between bits of genome and bits of phenotype, any more than there is mapping between crumbs of cake and words of recipe. The recipe is not a blueprint that can be reconstructed from

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. Theory 6. Direction (order) imposed on random variation by natural selection Darwinism – the non-random selection of randomly varying replication entities by reason of their ‘phenotypic’ effects – is the only force I know that can, in principle, guide evolution in the direction of adaptive complexity. It works on this planet. It

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efficacy throughout the universe. The ingredients in a general recipe for Darwinian evolution are replicating entities of some kind, exerting phenotypic ‘power’ of some kind over their replication success. In The Extended Phenotype I referred to these necessary entities as ‘active germ-line replicators’ or ‘optimons’. It is important to keep their

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replication conceptually separate from their phenotypic effects, even though, on some planets, there may be a blurring in practice. Phenotypic adaptations can be seen as tools of replicator propagation. Gould disparages the replicator’s-eye view of evolution as

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happy one: it is easy to see the genetic changes that accompany evolution as book-keeping entries, mere accountant’s records of the really interesting phenotypic events going on in the outside world. Deeper consideration, however, shows that the truth is almost the exact opposite. It is central and essential to

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Darwinian (as opposed to Lamarckian) evolution that there shall be causal arrows flowing from genotype to phenotype, but not in the reverse direction. Changes in gene frequencies are not passive book-keeping records of

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phenotypic changes: it is precisely because (and to the extent that) they actively cause phenotypic changes that evolution of the phenotype can occur. Serious errors flow, both from a failure to understand the importance of this one

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is, I believe, the force underlying all adaptive complexity. Other topics for a future science of Universal Darwinism Active germ-line replicators together with their phenotypic consequences, then, constitute the general recipe for life; but the form of the system may vary greatly from planet to planet, both with respect to

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the replicating entities themselves, and with respect to the ‘phenotypic’ means by which they ensure their survival. Indeed, as Leslie Orgel has pointed out to me, the very distinction between ‘genotype’ and

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‘phenotype’ may be blurred. The replicating entities do not have to be DNA or RNA. They do not have to be organic molecules at all. Even

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analogue, as opposed to digital codes, similar to the theoretical problems that would be raised by a purely analogue nervous system.*20 As for the phenotypic levers of power by which replicators influence their survival, we are so used to their being bound up into discrete organisms or ‘vehicles’ that we

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forget the possibility of a more diffuse extra-corporeal or ‘extended’ phenotype. Even on this Earth a large amount of interesting adaptation can be interpreted as part of the extended phenotype. There is, however, a general theoretical case that can be made in favour of the discrete

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with selection allows greater perfection of adaptation than selection on its own. An excellent and brilliant suggestion. *18 The one-way flow from genotype to phenotype – from genes to bodies – becomes obvious when you contrast the effect of a gene mutation (bodies change in future generations) with a purely bodily ‘mutation

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for treating the gene as special in this list of levels was clearly presented by Williams in 1966: The natural selection of phenotypes cannot in itself produce cumulative change, because phenotypes are extremely temporary manifestations. The same argument holds for genotypes…Socrates’ genes may be with us yet, but not his genotype

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neo-Darwinian ‘individual selection’. Nor did I, when I reiterated and extended the same argument a decade later in The Selfish Gene and The Extended Phenotype. We thought we were just clarifying what orthodox neo-Darwinism really meant. Yet both critics and supporters misunderstood our view as an attack on the

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of its class. An individual organism, such as a clonally reproducing aphid or stick insect, would be a true replicator only if blemishes in the phenotype – say an amputated leg – were reproduced in the next generation. And of course they are not. Note that a blemish in the genotype – a mutation

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– is reproduced in the next generation. Of course it may then show itself in the phenotype too, but it is not the phenotypic blemish itself which is copied. This is no more than the familiar principle of the non-inheritance of acquired characteristics, or – its

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, which implies that blemished replicators may be less proficient, or more proficient, than the original (in practice because of what we are accustomed to calling ‘phenotypic effects’). The true unit of selection in any Darwinian process, on any planet, is an active germ-line replicator. On this planet, it happens to

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plants ‘figure out’ the formula of chlorophyll? Enough, let us be constructive. Natural selection chooses genes rather than their alleles,*9 because of those genes’ phenotypic effects. In the case of behaviour, the genes presumably influence the state of the nervous system, which in turn influences the behaviour. Whether it is

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behaviour, physiology or anatomy, a complex phenotype may require sophisticated mathematical description if we are to understand it. This does not, of course, mean that the animals themselves have to be mathematicians

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results from a misunderstanding about what it means to speak of a gene ‘for’ behaviour. No geneticist has ever imagined that a gene ‘for’ some phenotypic character such as microcephaly, or brown eyes, is responsible, alone and unaided, for the manufacture of the organ that it affects. A microcephalic head is

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new gene of interest simply gave this existing complex process a crude kick, the end result of which was a crucial change in the complex phenotypic effect. What had been complex maternal care, say, became complex sibling care. The shift from maternal to sibling care was a simple one, even if

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creatures, natural selection can be seen as competition between alleles in the gene pool for that slot. The weapons of their competition are normally the ‘phenotypic’ effects that they have on bodies. *10 See also the footnote on this page. *11 ‘Evolutionarily stable strategy’ or ESS is John Maynard Smith’s

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in which a group phenomenon could be subject to genetic variation. You might contrive it via some version of what I have called the ‘extended phenotype’, but I am too sceptical to accompany you on that theoretical journey. My point, of course, is that the phenomenon of religion may be like

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is a gene: not the physical molecule of DNA but the information it carries. Biological codices, or genes, are carried around inside bodies whose qualities – phenotypes – they helped to influence. The death of the body entails the destruction of any codices that it contains, unless they have previously been passed on

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(London, Bantam, 2015) Dawkins, Richard, Climbing Mount Improbable (London, Viking, 1996) Dawkins, Richard, A Devil’s Chaplain (London, Weidenfeld & Nicolson, 2003) Dawkins, Richard, The Extended Phenotype (London, Oxford University Press, 1982) Dawkins, Richard, The God Delusion (London, Bantam, 2006; 10th anniversary edn, London, Black Swan, 2016) Dawkins, Richard, The Greatest Show

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Story of God: a personal journey into the world of science and religion (London, Bantam, 2005) Also by Richard Dawkins The Selfish Gene The Extended Phenotype The Blind Watchmaker River Out of Eden Climbing Mount Improbable Unweaving the Rainbow A Devil’s Chaplain The Ancestor’s Tale The God Delusion The

Blueprint: The Evolutionary Origins of a Good Society

by Nicholas A. Christakis  · 26 Mar 2019

that genes shape human behavior. The study of genetics and heredity began with how genotypes (genes and their variants) shape phenotypes (an organism’s physical appearance and function). But phenotypes eventually came to be seen as any manifestations of genes, moving beyond physical appearance to include the way the brain works

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analyzing tiny variations in DNA, behavior geneticists have explored whether the genome as a whole and also certain genes in particular help to explain complex phenotypes like neuroticism, decision-making, and friendliness. So broad and powerful is the impact of genes on behavior that, in 2000, psychologist Eric Turkheimer formulated

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people manifest numerous traits, from religiosity to risk aversion. Still, figuring out which genes really matter for a given complex behavioral trait (or any phenotype) can be difficult. It can be like trying to learn what makes a car run if you have never seen one before. You might discover

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is the “cause” of the car’s operation. There are many parts in a car that must work together to make it move. Some phenotypes are indeed simple, however. You might remember from high-school biology that different variants of the hemoglobin gene encode the production of different kinds of

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not like a simple computer program that runs exactly the same in every computer. There is rarely a one-to-one relationship between genotype and phenotype. Instead, genes yield particular behaviors more than others on average—just like smoking cigarettes substantially increases your chance of getting lung cancer on average

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another, a crucial idea regarding inter-individual genetic effects to which we shall return. There may also be associations between vasopressin-receptor expression and other phenotypes in humans as varied as age at first sexual intercourse, autism, and even, intriguingly, altruism.40 This further supports the idea that the evolution

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Ronald A. Fisher and Sewall Wright separately proposed the idea that if spouses resembled each other superficially (that is, phenotypically), they would also resemble each other genetically.47 Since most phenotypes arise from the actions of many genes working together, assortative (or disassortative) mating could result in genetic correlations between

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reared apart sometimes report an overwhelming (and usually terribly unwelcome) physical attraction when they meet as adults.86 In addition to location and individual identity, phenotype matching is another and even more complex possible mechanism for kin detection. Here, an organism evaluates how similar others are to itself. For instance,

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to have someone very different from you to help you with your math homework. Friends may resemble one another in subtle ways short of overt phenotypes. An important study by psychologist Thalia Wheatley and her colleagues Carolyn Parkinson and Adam Kleinbaum examined the brain responses of a group of forty-

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the pattern of blood flow in their brains in response to this sample of clips! Friends may resemble one another on a genotypic, not just phenotypic, level.32 There are four reasons for this. First, similar genotypes may simply reflect that friends often come from the same place. If Greeks

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when that ability is beneficial. The features animals use to identify others can be categorized into two types: cues and signals. Identity cues are phenotypic traits that make it possible to tell one individual from another but that do not themselves confer a survival advantage. In humans, fingerprints are unique

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.18 Fingerprints, like the unique pattern of small blood vessels in everyone’s eyes, are therefore merely a possible cue. Identity signals, however, are phenotypic traits that facilitate individual recognition while also assisting an animal’s survival. If you do not want others to mistakenly attack you, neglect to repay

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as their offspring, then you need some way to indicate This is me, not someone else.19 In order to achieve this, the relevant phenotypic trait should have a lot of variation so that it can be noticeably distinctive and memorable. As we would therefore expect, facial traits show more

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the details of human faces should be combinable into endless configurations (see figure 9.3).21 Variation in human faces is so important that rare phenotypes that help individuals appear special should be favored by natural selection. Consequently, there is indeed a lot of variation in the genes that code for

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be understood at many different levels, often reflecting scientists’ arbitrary interests. Biochemists might call it a day after studying how genes affect cells and observing phenotypes at the very first stage, in which genes are translated into their corresponding proteins. But why stop there? Medical geneticists may ignore the effect

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study how different genes affect muscle function or brain structure or disease symptoms. Zoologists interested in whole animals may do breeding manipulations to study a phenotype they are interested in, such as the coat color of foxes or the monogamy behavior of voles. Behavior geneticists may ignore these intermediate levels

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of social groups, an exophenotype.6 This type of idea was first advanced by evolutionary biologist Richard Dawkins in his profound 1982 book The Extended Phenotype. Dawkins made a case that it should theoretically be possible “to free the selfish gene from the individual organism which has been its conceptual

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perspective, a beaver may be wired to make a useful dam just as it is wired to have a functional pancreas. Dawkins speaks of extended phenotypes, but I prefer to use the term exophenotype, by which I mean the nonincidental, genetically guided changes that an organism makes to its surroundings

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, then these genes have a downstream effect on other people, and it might be this indirect effect outside the body that is making the phenotype advantageous. A skunk warding off predators or a flower attracting bees is changing the behavior of other organisms in order to improve its own fitness

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should encode the specific exophenotypes and that these encoded exophenotypes should affect reproduction or survival. Dawkins notes that the buildings people construct are not extended phenotypes, because humans do not, in fact, have genes that affect the propensity to build dwellings or genes that encode for whether those dwellings will

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in contact with. Third, and perhaps most remarkable, genes can influence organisms at a distance. The genotype of one living thing can affect the phenotype (including the behavior) of another member of its own species or of a different species without ever coming into physical contact with it. If human

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. Animal Artifacts Animal artifacts are material objects created by animals as a deliberate result of their actions; bird bowers are an example. Like any phenotype whose variation is influenced by a genotype, artifacts can help or hurt the chances that the organism will survive and reproduce and that the relevant

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a spider evolved to have bigger mouth parts at the front of its body to more easily catch prey, it would be seen as a phenotypic trait arising from genotypic change that affects its survival. But the web a spider places outside its body is really no different; by using

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on living tissue surrounding them rather than on inanimate matter. Snails infested with a certain kind of parasitic fluke have thicker shells. The shell phenotype is clearly influenced by the snail’s own physiology and genes. But the change in the host’s morphology may also be seen, at

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a parasite evolves to induce anatomical changes in its host in ways that enhance the parasite’s prospects for survival, we can say that the phenotype of the host is under the control of the genotype of the parasite. From the conventional point of view of the host, the modifications

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the latter into a spore-delivery platform. We know from fossilized leaves showing telltale marks of ant bites on their veins that this fungal phenotype is tens of millions of years old.28 Is it possible that some human traits and behaviors may actually be the genetic by-product of

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of my children observed, although in less clinical language).35 How can we focus our scientific gaze? A crucial consideration is this: for a phenotype to be relevant, it must result in the differential propagation of the underlying genes. Variants of genes can increase or decrease in frequency in a

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population as a consequence of their phenotypic effects, but some phenotypic effects may be incidental and not have any impact on the frequency of alleles in the next generation. To illustrate this point, consider

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for food and a safer route to swim from place to place. This pond can thus be seen as an exophenotype, different than bodily phenotypes like the size or shape of the beaver’s teeth. By modifying the size and depth of the pond, the beaver can create selection

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As a result, the capacity to build dams (an exophenotype) and the possession of bodily features that allow a beaver to swim far (a conventional phenotype) may coevolve. Regulating One Another’s Genes Something similar to beaver dams, I think, applies to the social structures humans create. If gene variants predispose

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can help us understand indirect genetic effects and hence such social, action-at-a-distance exophenotypes. The feather condition of a chicken is clearly a phenotype, with genes playing a role in a bird’s plumage. Environmental factors, including the chicken’s diet and light exposure, are relevant too, of

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going bald). If one gene changes, it may have an effect on the action of other genes, and hence on the expression of the attendant phenotypes. As we have just seen with the chickens, however, such effects can also occur between individuals. I call this social epistasis, since the impact

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especially if the groups espousing them are in competition with groups espousing less advantageous ideas. Culture can also manifest convergent evolution, just like genetically based phenotypes in animals, as we saw earlier. Consider the form of fishhooks, which have been independently invented by humans more than once. There are several

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a physical object made by an animal that has a structure specified by the animal’s genes. Plate 8: A “Zombie” Ant The extended phenotype of the fungus Ophiocordyceps unilateralis is manifested in the Asian ant Polyrhachis armata. The fungus controls the ant’s behavior, prompting it to climb a

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. 40. Z. M. Prichard, A. J. Mackinnon, A. F. Jorm, and S. Easteal, “AVPR1A and OXTR Polymorphisms Are Associated with Sexual and Reproductive Behavioral Phenotypes in Humans,” Human Mutation 28 (2007): 1150; T. H. Wassink et al., “Examination of AVPR1a as an Autism Susceptibility Gene,” Molecular Psychiatry 9 (2004): 968

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Ethology and Sociobiology 6 (1985): 183–187. 51. T. Antal, H. Ohtsuki, J. Wakeley, P. D. Taylor, and M. A. Nowak, “Evolution of Cooperation by Phenotypic Similarity,” PNAS: Proceedings of the National Academy of Sciences 106 (2009): 8597–8600; M. A. Nowak, “Five Rules for the Evolution of Cooperation,” Science 314

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al., “Effects of Experience on Fetal Voice Recognition,” Psychological Science 14 (2003): 220–224. 86. M. Greenberg and R. Littlewood, “Post-Adoption Incest and Phenotypic Matching: Experience, Personal Meanings and Biosocial Implications,” British Journal of Medical Psychology 68 (1995): 29–44. For a collection of reports of this phenomenon in

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Specialized Skills of Social Cognition: The Cultural Intelligence Hypothesis,” Science 317 (2007): 1360–1366. 41. Such an effect would especially speed up the evolution of phenotypes that are intrinsically synergistic, and this may help shed light on the observation that evolution in humans is accelerating. J. Hawks, E. T. Wang,

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Signatures of Convergent Evolution in Echolocating Mammals,” Nature 502 (2013): 228–231. In the case of echolocation, we know that genes used for the convergent phenotypes may be similar across widely divergent taxa, such as bats and dolphins. 9. S. C. Morris, Life’s Solution: Inevitable Humans in a Lonely

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Male Distinctiveness?,” Journal of Comparative Psychology 124 (2010): 343–350; C. Almstrom and M. Knight, “Using a Paired-Associate Learning Task to Assess Parent-Child Phenotypic Similarity,” Psychology Reports 97 (2005): 129–137. 31. K. McComb, C. Moss, S. M. Durant, L. Baker, and S. Sayialel, “Matriarchs as Repositories of

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behaviors. B. John and K. R. Lewis, “Chromosome Variability and Geographic Distribution in Insects,” Science 152 (1966): 711–721. 7. R. Dawkins, The Extended Phenotype: The Long Reach of the Gene (Oxford: W. H. Freeman, 1982), p. vi. 8. Writing in 1982, Dawkins noted that there was scant evidence for

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that you are not the first one to have a particular idea. 9. Ibid. Dawkins returned to this topic twenty years later. R. Dawkins, “Extended Phenotype—But Not Too Extended: A Reply to Laland, Turner, and Jablonka,” Biology and Philosophy 19 (2004): 377–396. 10. H. Eiberg et al., “

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study. K. Kleisner, L. Priplatova, P. Frost, and J. Flegr, “Trustworthy-Looking Face Meets Brown Eyes,” PLOS ONE 8 (2013): e53285. 16. Dawkins, “Extended Phenotype.” 17. Housing architecture belongs to the realm of culture; however, architecture might affect our evolution by other means, a process known as gene-culture coevolution

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, while species with colorful plumage tend to build less impressive bowers, as if some species over time have shifted from emphasizing physical phenotypes to emphasizing behavioral exophenotypes. Dawkins, Extended Phenotype, p. 199. 22. J. N. Weber, B. K. Peterson, and H. E. Hoekstra, “Discrete Genetic Modules Are Responsible for Complex

…

Burrow Evolution in Peromyscus Mice,” Nature 493 (2013): 402–405. 23. D. P. Hughes, “On the Origins of Parasite Extended Phenotypes,” Integrative and Comparative Biology 54 (2014): 210–217. 24. W. M. Ingram, L. M. Goodrich, E. A. Robey, and M. B. Eisen, “Mice Infected

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becoming more fertile in a way that was beneficial to the organism, then, yes, this would be seen as an exophenotypic effect. 36. Dawkins, Extended Phenotype, pp. 206–207. 37. L. Glowacki, A. Isakov, R. W. Wrangham, R. McDermott, J. H. Fowler, and N. A. Christakis, “Formation of Raiding Parties

…

, “Across-Line SNP Association Study for Direct and Associative Effect on Feather Damage in Laying Hens,” Behavior Genetics 40 (2010): 715–727. 39. Dawkins, Extended Phenotype, p. 230. 40. P. Lieberman, “The Evolution of Human Speech,” Current Anthropology 48 (2007): 39–66; D. Ploog, “The Neural Basis of Vocalization,” in

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2929–2933. 56. Y. Itan, B. L. Jones, C. J. E. Ingram, D. M. Swallow, and M. G. Thomas, “A Worldwide Correlation of Lactase Persistence Phenotype and Genotype,” BMC Evolutionary Biology 10 (2019): 36. 57. S. A. Tishkoff et al., “Convergent Adaptation of Human Lactase Persistence in Africa and Europe,” Nature

Epigenetics: How Environment Shapes Our Genes

by Richard C. Francis  · 14 May 2012

had to take a much more indirect route. He could only identify a gene through a mutation that caused some observable change in the appearance (phenotype) of his subjects. If this mutation was correlated with a different trait, he could assume that the genes for the two traits were located on

…

a result of epigenetic processes. The same may be true of obesity and its associated disorders in people like Paradorn. From Thrifty Genes to Thrifty Phenotypes Of course, our genes affect our weight. At issue is whether, in coming to grips with the obesity epidemic, our research dollars are best spent

…

Barker proposed that when the fetus receives insufficient nutrition through the placenta, it becomes programmed in the womb for a thrifty phenotype.9 As was proposed for the thrifty-genes hypothesis, those with a thrifty phenotype have a more efficient metabolism than babies born at a normal birth weight. But the thrifty

…

phenotype can result from diverse genetic backgrounds and without the aid of specific obesity genes. It is, rather, simply a function of the

…

intrauterine environment. The thrifty phenotype works out well in traditional non-Western cultures where the postnatal environment is often one of scarcity. In those cases, the prenatal environment predicts the

…

in an adaptive way. Problems arise, however, if the postnatal environment is enriched food-wise relative to the prenatal environment. When this mismatch occurs, thrifty phenotypes result in obesity and its consequences. The Barker hypothesis nicely accounts for the correlation between low birth weight and adult obesity and has been bolstered

…

? Barker himself has little interest in the mechanism through which this effect of the uterine environment occurs. Others, though, have pursued mechanistic investigations of thrifty phenotypes, and, as is usually the case, the initial studies have been conducted on nonhuman mammals—especially mice, rats, and sheep. For studies of this sort

…

color, and hence epigenetic state, of the mother. Female mice with the yellow coat color tend to produce yellow offspring, never offspring with the pseudoagouti phenotype. Mothers with the pseudoagouti coloration produce few yellow offspring and more pseudoagoutis.10 Moreover, a grandmother’s coloration also influences the coat coloration of her

…

The Mule Page, http://www.phudpucker.com/mules/mule.htm. 2. The Reivers (1962). 3. There is a characteristic cognitive deficit known as Turner neurocognitive phenotype (Ross, Roeltgen, et al. 2006), which is primarily restricted to spatial and mathematical reasoning. Turner syndrome is also associated with autism. 4. Parent-of-origin

…

passed into disuse, as a suitable name for the branch of biology which studies the causal interactions between genes and their products which bring the phenotype into being” (Waddington 1968). 30. In essence, Waddington’s goal was a synthesis of what was then known as “embryology” and genetics; we now call

…

). “Epigenetic mutations of the imprinted IGF2-H19 domain in Silver-Russell syndrome (SRS): Results from a large cohort of patients with SRS and SRS-like phenotypes.” J Med Genet 46(3): 192–197. Beadle, G. W., and E. L. Tatum (1941). “Genetic control of biochemical reactions in Neurospora.” Proc Natl Acad

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inactivation.” Trends Genet 19: 432–438. Burdge, G. C., M. A. Hanson, et al. (2007). “Epigenetic regulation of transcription: A mechanism for inducing variations in phenotype (fetal programming) by differences in nutrition during early life?” Br J Nutr 97(6): 1036–1046. Burmeister, S. S., V. Kailasanath, et al. (2007). “Social

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–617. Costa, F. F. (2008). “Non-coding RNAs, epigenetics and complexity.” Gene 410(1): 9–17. Coventry, W. L., S. E. Medland, et al. (2009). “Phenotypic and discordant-monozygotic analyses of stress and perceived social support as antecedents to or sequelae of risk for depression.” Twin Res Hum Genet 12(5

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., and D. A. Melton (2008). “Nuclear reprogramming in cells.” Science 322(5909): 1811–1815. Hales, C. N., and D. J. P. Barker (2001). “The thrifty phenotype hypothesis: Type 2 diabetes.” Br Med Bull 60(1): 5–20. Hamelin, C. E., G. Anglin, et al. (2006). “Genomic imprinting in Turner syndrome: Effects

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): 588–595. Hayashi, T., A. G. Motulsky, et al. (1999). “Position of a ‘green-red’ hybrid gene in the visual pigment array determines colour-vision phenotype.” Nat Genet 22(1): 90–93. Hayes, T. B., A. A. Stuart, et al. (2006). “Characterization of atrazine-induced gonadal malformations in African clawed frogs

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‘vitalism.’” Cell 100(1): 79–88. Ko, J. M., J. M. Kim, et al. (2010). “Influence of parental origin of the X chromosome on physical phenotypes and GH responsiveness of patients with Turner syndrome.” Clin Endocrinol (Oxf) 73(1): 66–71. Kochanska, G., R. A. Barry, et al. (2009). “Early attachment

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321(7276): 1609–1612. Kulesa, P. M., J. C. Kasemeier-Kulesa, et al. (2006). “Reprogramming metastatic melanoma cells to assume a neural crest cell-like phenotype in an embryonic microenvironment.” Proc Natl Acad Sci USA 103(10): 3752–3757. Lanctot, C., T. Cheutin, et al. (2007). “Dynamic genome architecture in the

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D. Ugarkovic (2008). “Role of non-coding RNA and heterochromatin in aneuploidy and cancer.” Semin Cancer Biol 18(2): 123–130. Pigliucci, M. (2010). “Genotype-phenotype mapping and the end of the ‘genes as blueprint’ metaphor.” Phil Trans Royal Soc B 365(1540): 557–566. Pollard, K. S., S. R. Salama

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.” Curr Opin Clin Nutr Metab Care 6(4): 369–375. Szyf, M., I. C. Weaver, et al. (2005). “Maternal programming of steroid receptor expression and phenotype through DNA methylation in the rat.” Front Neuroendocrinol 26(3–4): 139–162. Taft, R. J., M. Pheasant, et al. (2007). “The relationship between non

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radiata) and pigtail (Macaca nemestrina) macaques is related to early maternal experiences.” Am J Primatol 62(4): 243–259. Weaver, I. C. (2009). “Shaping adult phenotypes through early life environments.” Birth Defects Res C Embryo Today 87(4): 314–326. Weaver, I. C., N. Cervoni, et al. (2004). “Epigenetic programming by

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77(3): 151–158. Wolfram, S. (2002). A new kind of science. Champagn, IL: Wolfram Media. Wong, A. H., Gottesman, I. I., et al. (2005). “Phenotypic differences in genetically identical organisms: The epigenetic perspective.” Hum Mol Genet 14(Spec 1): R11–R18. Wright, S. (1916). “An intensive study of the inheritance

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Parent Bonding Instrument (PBI) parenting see also mothering parent-of-origin effect see also imprinting, genomic paternal care pathogen defense, growth vs. Petchburi Province, Thailand phenotypes thrifty pheromones pituitary placenta, imprinting in plants, epigenetic inheritance in pleiotropy pluripotent cells polychlorinated biphenyls (PCBs) population genetics postnatal environment, mismatch between fetal environment and

…

, Edward teratocarcinoma cells testes endocrine disruptors and steroid abuse and shrinkage of testosterone competitive interactions and Thailand, diet and obesity in “thrifty genes” hypothesis thrifty phenotype thymine tigons tissue-based theory of cancer Toguchi, Audrey Toronto zoo totipotent cells traits: genes as linked to sex-linked transcription transcription factors transdifferentiation transgenerational

…

epigenetic processes translation translocation tumor suppressor genes Turner neurocognitive phenotype Turner syndrome unipotent cells viable yellow (Avy) agouti allele Vietnam Veterans Memorial Vietnam War vinclozolin Vindicated (Canseco) violence, steroid use and vision, color, see color

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