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,

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

Darwin's Dangerous Idea: Evolution and the Meanings of Life

by Daniel C. Dennett  · 15 Jan 1995  · 846pp  · 232,630 words

or revise their wiring, depending on what they encounter during their lifetimes. (In the language of evolutionary theory, there is some "plasticity" in their phenotypes. The phenotype is the eventual body design created by the genotype in interaction with environment. Identical twins raised in different environments would share a genotype but might

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be dramatically different in phenotype.) Suppose, then, that these organisms can end up, after exploration, with a design different from the one they were born with. We may suppose

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good that those who never learn it, or who learn it "too late," are at a severe disadvantage. In populations with this sort of phenotypic plasticity, a near-miss is better than a mile. For such a population, the telephone pole in the desert becomes the summit of a gradual

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genotype level — will tend to follow the lead o/and confirm the directions taken by the individual organisms' successful explorations — redesign at the individual or phenotype level. The way I have just described the Baldwin Effect certainly keeps Mind to {79} FIGURE 3.2 a minimum, if not altogether out of

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would not usefully be regarded as a gene for not reading. This is simply because preventing reading would not be its most obvious or debilitating phenotypic effect. [Dawkins 1982, p. 23. See also Dawkins 1989a, pp. 281-82, and Sterelny and Kitcher 1988.] The indirect way in which groups of

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of Biomorph Land, Dawkins stresses that a tiny — indeed minimal — change in the genotype (the recipe) can produce a strikingly large change in the phenotype (the resulting individual organism), but he tends to slight one of the major implications of this: if a single step in the genotype can produce

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a giant step in the phenotype, intermediate steps for the phenotype may be simply unavailable, given the mapping rules. To take a deliberately extreme and fanciful example, you might think that if a

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concept of biological possibility: x is biologically possible if and only if x is an instantiation of an accessible genome or a feature of its phenotypic products. Accessible from where? By what processes? Ah, there's the rub. We have to specify a starting point in the Library of Mendel,

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to have a fresh turkey dinner on Christmas Day, 2001, if and only if at least one instantiated turkey genome has produced the requisite phenotypic effects in time for dinner. It is biologically possible for you to ride a pteranodon before you die if and only if Jurassic Parkish technology

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stopped meandering about sixty million years ago.) What rules govern travel through this space? What rules or laws constrain the relations between genomes and their phenotypic products? So far, all we have acknowledged are logical or mathematical necessities on the one hand and the laws of physics on the other. That

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of "irreducible biological law." For instance, many have argued that there are "developmental laws" or "laws of form" that constrain the relation between genotype and phenotype. In due course we will consider their status, but already we can locate at least some of the most salient constraints on biological possibility as

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are, and so are the genomes of all the life forms our lives depend on). The Library of Babel describes one aspect of our "extended phenotype" (Dawkins 1982). That is, in the same way that spiders make webs and beavers make dams, we make (among many other things) books. You

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to wander into the attractive basin of a new norm.10 Reproductive isolation is thus both a cause and an effect of the clumpiness of phenotypic space. Wherever there are competing error-correcting regimes, one regime or the other will win out, and hence the isthmus between the competitors will

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much more difficult to get any epistemic purchase on the developmental or building {217} process by which a genotype gets "expressed" in a fully formed phenotype. And the design processes that shaped the developmental processes that shape the "finished products" were largely inaccessible to the sorts of intrusive observation and manipulation

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and hence von Baer's laws, do not represent a special mechanism of developmental canalization, the usual sense of which is a buffering of the phenotype against genetic alteration... Instead, locking-in of early development is a direct reflection of the fact that the number of ways to improve organisms by

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"Darn — organisms are 'too smart' to cooperate!"16) The standard assumption of game theory is that there will always be mutations that have the "right" phenotypic effects to rise to the occasion, but what if the right move just doesn't "occur to Mother Nature"? Is this ever or often very

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other direction, supposing that the space of real possibilities is much more densely populated than it actually is. The temptation, when we think about phenotypic variation, is to adopt a sort of Identikit tactic of assuming that all the minor variations we can imagine on the themes we find in

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some rather bushier but still far from dense partial filling of the apparently possible. We have already seen that the Vast space of all imaginable phenotypes — Identikit Space, we might call it — no doubt includes huge regions for which there are no recipes in the Library of Mendel. But even

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often illustrated by contrasting a pair of trees of life (figure 10.6). We can suppose that the horizontal dimension registers some one aspect of phenotypic variation or body design — we'd need a multidimensional space to represent it all, of course. The orthodox view on the left is pictured

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it needs more dimensions, so we can compare the steps in genotype space (the typographical steps in the Library of Mendel) to the steps in phenotype space (the design innovations in Design Space) and then evaluate these differences on a fitness landscape. As we have seen, the relations between recipe

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three different reasons we could call a mutation big: it is a big step in the Library of Mendel; it produces a radical difference in phenotype (a monster); it produces (one way or another) a big increase in fitness — a lot of lifting, in our metaphor for good work done

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in the large portion of DNA that never gets expressed, and if these accumulated changes suddenly got expressed, thanks to some transposing error, a huge phenotypic effect would be expected. But it is only when we turn to the third sense of macromutation — large differences in fitness — that we can

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of Goldschmidt's notorious book, he expanded on this point: Darwinians, with their traditional preferences for gradualism and continuity, might not shout hosannas for large phenotypic shifts induced rapidly by small genetic changes that affect early development; but nothing in Darwinian theory precludes such events, for the underlying continuity of small

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describe this possibly underappreciated fact about development as a non-Darwinian creative force in evolution, "because the constraints that it imposes upon the nature of phenotypic change guarantee that small and continuous Darwinian variation is not the raw material of all evolution," for it "relegates selection to a negative role (eliminating

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a king would be unnerving, but what if no such deliberate clues were left around? Would a closer look at the organism designs themselves — the phenotypes — reveal some telltale discontinuities? Gene-splicers {317} are the most powerful cranes we have yet discovered. Are there designs that simply could not be

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is the actual "decision-making" of natural selection most directly responsive? It is not controversial that conflicts between genes and bodies (between genes and the phenotypic expressions of the genotypes of which they are a proper part) can arise. Moreover, no one doubts that in general the body's claim to

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produce such a big effect. What have changed dramatically are human health, diet, and living conditions; these are what have produced the dramatic change in phenotype, which is 100 percent due to cultural innovations, passed on through cultural transmission: schooling, the spread of new farming practices, public-health measures, and

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be due to cultural changes, since fewer than two hundred generations separate us from Plato. Environmental changes due to cultural innovations change the landscape of phenotypic expression so much, and so fast, however, that they can in principle change the genetic selection pressures rapidly — the Baldwin Effect is a simple

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are clearly a meme, replicating all by themselves, detached from the rest of the symphony, but keeping intact a certain identity of effect (a phenotypic effect), and hence thriving in contexts in which Beethoven and his works are unknown. Dawkins explains how he coined the name he gave these units

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turn out that cultural evolution recapitulates all the features of genetic evolution: not only are there gene analogues (memes), but there are strict analogues of phenotypes, genotypes, sexual reproduction, sexual selection, DNA, RNA, codons, allopatric speciation, demes, genomic imprinting, and so forth — the whole edifice of biological theory perfectly mirrored

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, and are proving to be virtually unquarantinable. Genes are invisible; they are carried by gene vehicles (organisms) in which they tend to produce characteristic effects (phenotypic effects) by which their fates are, in the long run, determined. Memes are also invisible, and are carried by meme vehicles — pictures, books, sayings (

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, a meme's prospects depend on its design — not its "internal" design, whatever that might be, but the design it shows the world, its phenotype, the way it affects things in its environment. The things in its environment are minds and other memes. For instance, whatever virtues (from our perspective

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) the following memes have, they have in common the property of having phenotypic expressions that tend to make their own replication more likely by disabling or preempting the environmental forces that would tend to extinguish them: the meme

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course not — that's how powerful the conspiracy is!" Some of these memes are "good," and others "bad"; what they have in common is a phenotypic effect that systematically tends to disable the selective forces arrayed against them. Other things being equal, memetics predicts that conspiracy-theory memes will persist quite

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locked to the melody, virtually guarantees a chain of thought in most listeners that would be undesirable on almost any festive occasion. This is the phenotypic effect that prevents the greater replication of this meme. If performances of The Mikado waned over the years, so that a time came when

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as playing a major role in eye development in the homeobox of mice has almost the same codon spelling as a gene dubbed (for its phenotypic effect) eyeless when it was identified in the homeobox of the fruitfly, Drosophila. But we would be even more flabbergasted were we to discover

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film and other media!) Our interest in identifying and re-identifying genes over the evolutionary ages is similarly primarily because of the uniformity of the phenotypic effects — what they are "about" (such as making hemoglobin, or eyes). Our ability to rely on their syntactic identifiability in DNA is a recent

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The very creativity and activity of human minds as temporary homes for memes seems to guarantee that lines of descent are hopelessly muddled, and that phenotypes (the "body designs" of memes) change so fast that there's no keeping track of the "natural kinds." Recall, from chapter {356} 10 (p.

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abstract formulation that is predictive precisely because it ignores all those complications. For instance, genes are selected because of their indirect and only statistically visible phenotypic effects. Consider the following prediction: wherever you find moths with camouflage on their wings, you will find that they have keen-sighted predators, and

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." Why did he retreat like this? Why, indeed, is the meme meme so little discussed eighteen years after The Selfish Gene appeared? In The Extended Phenotype (1982, p. 112), Dawkins replied forcefully to the storm of criticism from sociobiologists and others, while conceding some interesting disanalogies between genes and memes: ...

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, the spider's legs, the bird's wing. From this perspective, the vast protective networks of memes that we spin is as integral to our phenotypes — to explaining our competences, our chances, our vicissitudes — as anything in our more narrowly biological endowment. (This claim is developed in greater detail in

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many millions of cycles, producing many wonderful designs, both plant and animal, and eventually among its novel creations were some designs with the property of phenotypic plasticity. The individual candidate organisms were not wholly designed at birth, or, in other words, there were elements of their design that could be adjusted

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from the outside environment, via experience. Adaptationism is the view that the selecting environment gradually shapes the genotypes of organisms, molding them so that the phenotypes they command are some near-optimal fit with the encountered world. Behaviorism is the view that what Skinner (1953, especially pp. 129-41) called "

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belong to two very different categories. An average human is normally host to billions of symbiotic organisms belonging to perhaps a thousand different species.... His phenotype is not determined by his human genes alone but also by the genes of all the symbionts he happens to be infected with. The symbiont

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havens for endangered species, but at least they are havens, and what they preserve is irreplaceable. The same is true of complex memes and their phenotypic expressions. Many a fine New England church, costly to maintain, is in danger of destruction. Shall we deconsecrate these churches and turn them into museums

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(1990), is responsible for the perspective that allows us to see species as variably capable of "seeing" design improvements, thanks to their variable capacities for phenotypic exploration (for commentary, see Dennett 1990a). 7. Robert Richards' account of the history of the Baldwin Effect (1987, especially pp. 480-503 and discussion

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According to this complaint, orthodoxy vastly overestimates the extent to which the DNA can be considered to be a recipe, composed of genes, specifying a phenotype or an organism. Those who make this claim are the deconstructionists of biology, elevating the reader to power by demoting the text. It is a

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their host organisms, so we shouldn't get our hopes up unduly. See Engels 1992. 7. "I confess that I believe the emptiness of phenotypic space is filled with red herrings. ... Under the null hypothesis that no constraints at all exist, the branching pathways through space taken by this process

A Devil's Chaplain: Selected Writings

by Richard Dawkins  · 1 Jan 2004  · 460pp  · 107,712 words

English Edition, her interests lie in the conceptual mapping of knowledge and the thoughtful dissemination of learning. 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

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of adaptive evolution, but it assumes that only a minute fraction of DNA changes in evolution are adaptive in nature, while the great majority of phenotypically silent molecular substitutions exert no significant influence on survival and reproduction and drift randomly through the species. The facts of adaptation compel us to the

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information content in the Shannon sense. The best we can do is probably to forget about the genome itself and look at its product, the ‘phenotype’, the working body of the animal or plant itself. In 1951, J. W. S. Pringle, who later became my Professor at Oxford, suggested using a

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, docudrama (or docudrama) 2848, sociobiology 6679, catastrophe theory 1472, edge of chaos 2673, wannabee 2650, zippergate 1752, studmuffin 776, post-structural (or poststructural) 577, extended phenotype 515, exaptation 307. Of the 5042 mentions of memetic, more than 90 per cent make no mention of the origin of the word, which suggests

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experiments, a child somewhere along the line will forget some crucial step in the skill taught him by the previous child, and the line of phenotypes will suffer an abrupt macromutation which will presumably then be copied to the end of the line, or until another discrete mistake is made. The

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Lamarckian (Susan Blackmore calls it ‘copying the product’). In the origami experiment it is Weismannian (Blackmore’s ‘copying the instructions’). In the drawing experiment, the phenotype in every generation is also the genotype – it is what is passed on to the next generation. In the origami experiment, what passes to the

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a set of instructions for making it. Imperfections in the execution of the instructions result in imperfect junks (phenotypes) but they are not passed on to future generations: they are non-memetic. Here are the first five instructions in the Weismannian meme-line of

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the Chinese junk meme is made of paper. With the exception of ‘extended phenotypes’ such as beaver dams and caddis larva houses, the phenotypes of genes are normally parts of living bodies. Meme phenotypes seldom are. But it can happen. To return to my school again, a Martian geneticist, visiting the school during

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for many generations. Our Martian geneticist would have to work quite hard to discover that no genes are involved in the genesis of the roundhead phenotype. The Martian geneticist’s eyes would also pop out on stalks (assuming they weren’t on stalks to begin with) at the contemplation of certain

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styles of clothing and hairdressing, and their inheritance patterns. The black skullcapped phenotype shows a marked tendency towards longitudinal transmission from father to son (or it may be from maternal grandfather to grandson), and there is clear linkage

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to the rarer pigtail-plaited sideburn phenotype. Behavioural phenotypes such as genuflecting in front of crosses, and facing east to kneel five times per day, are inherited longitudinally too, and are in strong

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linkage disequilibrium with the previously mentioned phenotypes, as is the red-dot-on-forehead phenotype, and the saffron robes/shaven head linkage group. Genes are accurately copied and transmitted from body to body, but some are

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kind of neutral mutation will forever be hidden from the field biologist and from natural selection. And if a field biologist actually sees variation in phenotypes, the question of whether that variation could be selectively neutral cannot be settled in the biochemistry laboratory. Several essays touch on aspects of the relationship

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(s). Irrevocable determination by genes is no part of the idea nor is anything remotely approaching a ‘one gene, one trait’ mapping from genotype to phenotype. In any case it has nothing to do with ‘supreme confidence in universal adaptation’, which is as likely to be found among devotees of ‘individual

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metaphor, because it reverses the causal arrow, almost in Lamarckian fashion, and makes the genes passive recorders. I dealt with this in 1982 (The Extended Phenotype) in my distinction between ‘active replicators’ and ‘passive replicators’. The point is also explained in David Barash’s superb review of Gould’s book.136

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: Diversity, Evolution, and Inheritance (Cambridge, Mass., Harvard University Press, 1982) 65 F. H. C. Crick, Life itself (London, Macdonald, 1982) 66 R. Dawkins, The Extended Phenotype (San Francisco, W. H. Freeman, 1982/ Oxford, Oxford University Press, 1999), pp. 174–6. See also Endnote 36 and Dawkins, The Blind Watchmaker, chapter 11

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(London, Methuen, 1972) 117 R. Dawkins, The Selfish Gene, 2nd edn (Oxford, Oxford University Press, 1989), pp. 271–2. See also R. Dawkins, The Extended Phenotype (Oxford University Press, 1999), pp. 116–17, 239–47 118 Review of S. J. Gould, Wonderful Life (London, Hutchinson Radius, 1989). Published in the Sunday

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, (i) Evolutionarily stable state, (i) Exam Pressure, Destructive effects of, (i), (ii) Syllabuses, Limited nature of, (i) Expression of the Emotions, The, (i), (ii) Extended phenotype, (i), (ii) Extinction, (i), (ii), (iii), (iv), (v), (vi), (vii), (viii) Fabre, Jean Henri, (i) Faith As symptom of infection by mind virus, (i) Exercised

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) Theory of sex, (i) Parental expenditure/investment, (i), (ii) Patai, Daphne, (i) Pater, Walter, (i) Patriotism, Mindless, (i) Peacocks’ tails, (i) (see also Sexual selection) Phenotype, (i), (ii), (iii), (iv), (v), (vi) Phylogenetic tree, (i), (ii) Physics Envy, (i) Picasso, Pablo, (i) Pierce, Naomi, (i) Piezoelectric effect, (i) Piltdown hoax, (i

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

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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

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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

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

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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

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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

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? 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

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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

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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

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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

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). “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

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, 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

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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

The Barbell Prescription: Strength Training for Life After 40

by Jonathon Sullivan and Andy Baker  · 2 Dec 2016  · 742pp  · 166,595 words

Nassim Nicholas Taleb Introduction: Resistance is Not Futile Part I: Why – From Exercise Prescription to Training Program Jonathon M. Sullivan Chapter 1 The Sick Aging Phenotype Chapter 2 Exercise Medicine Chapter 3 From Prescription to Program: Safety and Dosing Chapter 4 Enduring Resistance, Resisting Endurance: Comprehensive Training Chapter 5 Specificity and

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Acknowledgements Notes Bibliography Authors Figures & Tables Chapter 1 Figure 1-1 Insulin signaling in health and disease Figure 1-2 Development of the Sick Aging Phenotype Chapter 4 Figure 4-1 The AMPK-Akt switch Figure 4-2 ATP – Adenosine triphosphate Figure 4-3 Energy transfer by ATP Figure 4-

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Table 4-2 Summary of the effect of different training approaches on energy systems and athletic attributes Chapter 5 Figure 5-1 The Sick Aging Phenotype (Reprise) Figure 5-2 Training increases both insulin-independent and insulin-dependent glucose flux in skeletal muscle Table 5-1 Maximal oxygen uptake (VO2max

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“seniors” the world has ever seen, or a ghastly and increasingly common syndrome of maladaptive aging, which we shall call the Sick Aging Phenotype. The Sick Aging Phenotype is a complex of interrelated and synergistic processes, in which the metabolic syndrome, muscle and bone loss, frailty, loss of function and independence

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instructions (genes) encoded in its DNA. Two organisms of the same species with identical or nearly identical genotypes will tend to have very similar phenotypes. But their phenotypes can also be very different. Allow me to give you an example. Consider a pair of identical twins, Will and Phil. Will and

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Easter, and they even seem to have similar temperaments. In other words, during development, childhood, and adolescence, Will and Phil demonstrate astonishingly similar phenotypes. Because their phenotypes are heavily influenced by their genotypes, this is not surprising. It is the genes, after all, that provide the source code for our biological

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$185,000 bill, which means Will can afford to go hiking in Scotland some 30 years later. Both Wellness Will and Phat Phil are modern phenotypes. In ages past, war, famine and infectious diseases were the scourge of mankind. Smallpox, diphtheria, cholera, measles, dysentery, plague, malaria, influenza, pneumonia, meningitis, cellulitis

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leading causes of mortality in industrialized societies,3 although a few idiot busybodies are working hard to undo these advances.4 Modern aging and death phenotypes are increasingly the product of abundance, longevity and idleness, with the major cardiovascular diseases (including stroke) being by far the number one cause

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remains considerable controversy about the particulars, especially with regard to the role of cultural influences, medical interventions, and diet.10 It appears the modern aging phenotypes are profoundly sensitive to a number of external and behavioral variables. Obviously, this book is focused on one of those variables: physical exercise. But

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hyperglycemia, and thence to full-blown diabetes. The other consequences are less obvious, but no less devastating. Figure 1-2. Development of the Sick Aging Phenotype. Abnormally high energy balance from diet and lack of exercise, along with genetic and other factors (top register), contribute to the development of obesity,

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“sick fat,” insulin resistance and metabolic syndrome. As this aging phenotype progresses (middle register), its components become increasingly intertwined and synergistic, culminating in a recalcitrant end stage (bottom register) characterized by full-blown diabetes, cardiovascular disease

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an increase in systemic inflammation. Inflammation promotes the development of atherosclerosis and degenerative changes in many tissues,33 promoting the development of the Sick Aging Phenotype. One of the principle sources of inflammatory products in the setting of obesity and metabolic syndrome is the fat itself. We now know that

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to the development of vascular disease, but not simply because serum triglycerides and cholesterol levels are elevated. Biological systems are complicated, and any particular phenotype is always the result of multiple factors. So it seems far more likely that vascular disease results from several primary effects of the metabolic syndrome

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induced oxidative stress promotes healthy biochemical adaptations that increase cellular tolerance to free radicals.19 Neurological health The depredations of aging and the Sick Aging Phenotype on the body are awful enough, but loss of neurological and cognitive functions are the cruelest cuts of all. This is an area where

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of the aging adult. * * * Requirements for the Exercise Prescription We’ve seen how a sedentary lifestyle contributes directly and substantially to the Sick Aging Phenotype and how exercise is a powerful medicine against the development of this slow-motion catastrophe. Of course, you already know this. You may not have

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must be comprehensive. Our exercise prescription should be as integrated and complete as possible. Our exercise prescription must specifically and effectively combat the Sick Aging Phenotype: It should attack the metabolic syndrome, reduce visceral fat, arrest or reverse sarcopenia and osteopenia, and fight frailty by retaining or restoring strength, power,

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to optimize performance in practice and play. In this chapter, we will see that programmed strength training effectively combats the components of the Sick Aging Phenotype, while allowing the Masters Athlete to optimize the General Fitness Attributes and retain muscle, bone, strength, power, and function. * * * Phat Phil Rises Again: Strength

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. In this chapter, we will look at the 4th of our criteria: Our exercise prescription/training program must specifically and effectively combat the Sick Aging Phenotype: metabolic syndrome, cardiovascular disease, sarcopenia, osteopenia, and loss of strength, function and mobility. After all, we’ve decided that the “event” we’re

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), and his tendons got floppier, weaker, and more easily injured. Phil developed frailty, chronic pain, depression, polypharmacy and loss of functioning. His Sick Aging Phenotype ultimately killed him through coronary artery disease and a massive heart attack, but it could just as easily have done him in with congestive heart

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of the metabolic syndrome promotes ongoing physiological imbalances, inflammation, and damage to tissue, especially vascular tissues. These processes culminate in the full-blown Sick Aging Phenotype: diabetes, atherosclerosis and vascular disease, heart disease, stroke, sarcopenia, weakness, frailty, chronic pain, and tissue atrophy. As the meta-syndrome develops, its component processes

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become more interlinked and reinforce each other, so the Sick Aging Phenotype becomes progressively more established and difficult to treat. Strength Training, Sick Fat, and Inflammation An abnormally positive energy balance is a principal cause of the

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strength training has the power to fight the accumulation of sick fat, a major player in the development of metabolic syndrome and the Sick Aging Phenotype.10 Strength Training, Metabolic Syndrome, and Diabetes Any form of regular, vigorous exercise combats the development and progression of the metabolic syndrome, and this

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insulin-dependent period.18 In other words, exercise promotes both insulin-independent and insulin-dependent glucose disposal,19 with profound effects on an insulin-resistant phenotype. The sheer volume of research data indicating that strength training combats insulin resistance and the metabolic syndrome has become overwhelming. Study after study shows that

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are to display the hellish constellation of visceral obesity, insulin resistance, hypertension, dyslipidemia, and systemic inflammation that points the way to the Sick Aging Phenotype. Accumulating evidence of the effect of resistance training on the metabolic syndrome was analyzed in depth by Strasser, Siebert, and Schobersberger in a study published

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development and management of Type 2 diabetes. Because of the centrality of insulin resistance to the development of the metabolic syndrome and the Sick Aging Phenotype, it is difficult to exaggerate the importance to the Masters Athlete of training for strength. Cardiovascular Health Because insulin resistance and the metabolic syndrome

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too old to do anything about it. Where are the Cheetos?” I believe that intrinsic human apoptosis is a powerful contributor to the Sick Aging Phenotype: a living hell of progressive weakness, obesity, inactivity, shrinking horizons, sexual impotence, decreased expectations, mounting despair, a growing list of expensive drugs, learned helplessness,

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Attributes STRENGTH LSD AEROBIC HIIT STRENGTH + LSD STRENGTH + HIIT Endurance ✔ ✔✔✔ ✔✔ ✔✔✔ ✔✔✔ Mobility and Balance ✔✔✔ No ✔ ✔✔✔ ✔✔✔ Strength ✔✔✔ No No ✔✔✔ ✔✔✔ Power ✔✔✔ No ✔ ✔✔✔ ✔✔✔ Body Composition ✔✔ ✔✔ ✔✔ ✔✔✔ ✔✔✔ Tier 3: Sick Aging Phenotype STRENGTH LSD AEROBIC HIIT STRENGTH + LSD STRENGTH + HIIT Physical inactivity and positive energy balance ✔✔✔ ✔✔✔ ✔✔✔ ✔✔✔ ✔✔✔ Obesity and visceral fat ✔✔ ✔✔✔ ✔✔✔ ✔✔✔ ✔✔✔ Insulin Resistance ✔✔ ✔✔✔ ✔✔✔ ✔✔✔ ✔✔✔ Cardiovascular Disease ✔ ✔✔✔ ✔✔✔ ✔✔✔ ✔✔✔ Sarcopenia ✔✔✔ No ✔ ✔✔ ✔✔✔

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shortcomings in the exercise medicine literature, an overwhelming amount of data clearly indicates that resistance training has profound and beneficial impacts on the Sick Aging Phenotype. How much more effective can we expect it to be when the medicine is prescribed and administered correctly? Your Physiological 401k When you’re

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for serious barbell training. Chapter 6 Simplicity and Efficiency: From Black Iron to Grey Steel Chapter Summary: The biology of aging, the Sick Aging Phenotype, and the effect of various exercise interventions on the physiology of aging populations indicate that strength training is the correct focus of a General Exercise

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attributes, and spans all three energy systems. And as we saw in the last chapter, a strength and conditioning program specifically addresses the Sick Aging Phenotype, combating insulin resistance, cardiovascular disease, sarcopenia, osteopenia, loss of mobility and function, frailty, depression, and the whole sick spiral of human apoptosis. Strength training

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, some degree of heart failure, brittle bones, threadbare muscles, a shopping bag full of prescription medicines, exercise intolerance, frailty, and loss of function. The phenotype comes to lethal harvest with stroke, heart attack, congestive heart failure, hip fracture, vertebral collapse, end-stage kidney disease, retinal hemorrhage or degeneration, toxic polypharmacy

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, and it is the back-strengthening exercise par excellence. The deadlift is safe, comprehensive, and has a wide therapeutic window. It fights the Sick Aging Phenotype, and is learned quickly by most athletes. This chapter covers the essentials of performance of the deadlift and discusses remedial variations. * * * What is

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relying entirely on the phosphagen system for the development of power. They emphasize grace, balance, mobility, and timing. The olympic movements attack the sick aging phenotype If they can be trained productively, the Olympic movements will confer beneficial impacts on glucose disposal, insulin sensitivity, neuromuscular and bioenergetic adaptation, cardiovascular health, frailty

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a broad therapeutic window. Our conditioning component must be as comprehensive as possible. Our conditioning component must specifically and effectively combat the Sick Aging Phenotype. It should also be as specific as possible to the physical demands confronted by the individual athlete. Our conditioning component should be as simple

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density, or mobility. But the addition of a conditioning component to our barbell prescription enhances the ability of strength training to fight the Sick Aging Phenotype, and our conditioning modality must fulfill this criterion. This is not a problem, because almost any type of exercise will improve cardiovascular and metabolic

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health. Decades of research have established that virtually all forms of conditioning have some activity against the Sick Aging Phenotype, reducing visceral fat, improving insulin sensitivity, and resisting the development and progression of cardiovascular disease. However, conditioning work also has implications for sport- and

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low-volume, high-intensity conditioning program, sometimes called a GPP (general physical preparedness program). Such a program specifically and effectively addresses the Sick Aging Phenotype while also fulfilling the requirement of a comprehensive preparation for a broad range of power-capacity demands. Conditioning simplicity Coaches and trainees often get too

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forms of conditioning work, running and jogging clearly confer cardiovascular benefits and physical performance improvements, and so they are partially effective against the Sick Aging Phenotype. However, their tendency to interfere with the far more comprehensive and important activity of strength training against unhealthy aging limits their fulfillment of this

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training stress, and increase the volume or frequency of deadlifts. Frequently voiced concerns that barbell training will endow the female Master with a “bulky,” masculine phenotype and grotesque muscular hypertrophy are so completely unfounded as to be ridiculous. In this chapter, therefore, we ridicule them. * * * Last But Not Least

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to increased mitochondrial biogenesis and improved oxidative efficiency, but more muscle protein breakdown. Each pathway inhibits the other, and concurrent training favors a more “aerobic” phenotype over strength. Anabolism: The part of metabolism that results in the accumulation of larger molecules and tissue mass. Contrast with catabolism. Anabolic Resistance: A

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population by virtue of meeting the criteria of safety, wide therapeutic window, comprehensive impacts on fitness attributes, specificity to and effectiveness against the Sick Aging Phenotype, simplicity and efficiency. General Fitness Attributes: The physical characteristics that positively impact the capacity and readiness of the organism to meet the physical demands

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programs in this book prescribe sets across – for example, the athlete will complete all squat sets before moving on to the press, etc. Sick Aging Phenotype: A meta-syndrome of pathological aging characterized by metabolic syndrome, sarcopenia and osteopenia, frailty, and polypharmacy. Detailed in Chapter 1. Sick Fat: See Visceral

Evil Genes: Why Rome Fell, Hitler Rose, Enron Failed, and My Sister Stole My Mother's Boyfriend

by Barbara Oakley Phd  · 20 Oct 2008

is intricately involved in the regulation of which genes are turned on or off in which tissues. This in turn ultimately determines an organism's phenotype—how an organism appears. Thus, although genes are still of paramount importance, there is a whole additional layer of complexity involving control of those genes

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contained on the long strands of DNA molecules in the nucleus of our cells—we can determine our genotype only by using molecular methods. Our phenotype, on the other hand, relates to our appearance, which is determined by the output of the genotype and sometimes by environment as well. Now we

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founding of orders and classes, is called macroevolution. Fig. 3.3. MACHIAVELLIAN GENES A complex array of varying genes underlies the many different outward manifestations—phenotypes—of many different personality disorders. A person with an unlucky shake of the genetic dice can actually end up with full-blown versions of those

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a personality disorder, he or she can be pushed into a full-blown version. Intermediate Phenotype Intermediate phenotype is a concept used by researchers who are wrestling with the relationship between genes and phenotype. To understand “intermediate phenotype,” it's helpful to remember that there is often an intermediate case between a full-blown

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manifestation of a disease and a less harmful variant. In personality disorders, intermediate phenotypes, sometimes called endophenotypes, are used to describe people with subclinical symptoms of diseases like schizophrenia or borderline personality disorder. The stipulation for an intermediate

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phenotype is that it be found in mildly ill but not “certifiable” siblings and other relatives, and that it even be found in some psychiatrically well

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related to risk for an illness and are not the illness itself. So far, the concept of intermediate phenotypes has been most powerfully developed and used by Dr. Michael Egan and his colleagues at the National Institute of Mental Health for their research on

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particular allele related to cognitive function that had previously been weakly and inconsistently associated with schizophrenia. When Egan's group applied the concepts of intermediate phenotype by studying brain function and comparing genotypes in a wide variety of people—including patients with schizophrenia, their healthy siblings, and controls—the suspect allele

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every person sampled, whether or not the person had schizophrenia. Egan's study was one of the first times that a correlation of an intermediate phenotype with a gene was shown to clarify how a gene related to a complex clinical diagnosis.9 “What's so surprising,” marvels Daniel Weinberger, Egan

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, “is that it works.”10 It appears that the next step beyond imaging genetics may relate to the synergistic use of genotyping, neuroimaging, and intermediate phenotypes. It will be exciting to see what future studies reveal when these techniques are applied to antisocial personality disorder, other related syndromes, and their subclinical

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“intermediate phenotypes.” Faced with the overwhelming variety of phenotypes that can arise from this mixture of genes and environment, it's hard to know where to even begin looking at a

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discipline, imaging genetics, has recently arisen that provides precisely the necessary tool. Imaging genetics uses medical imaging techniques to figure out a person's phenotype—the word phenotype meaning, in this case, the size and shape of organs such as the amygdala and cingulate cortex—and then evaluating the same person's

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our purposes, regarding traits such as impulsivity, mood, and anxiety. Through the use of such sophisticated new techniques and concepts as imaging genetics and intermediate phenotypes, researchers are discovering how alleles of particular genes can help underpin the dysfunctional behavior that can lead to a problematic personality or full-blown clinical

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narcissistic personality disorders).38 Differences in neural behavior in those with different versions of the gene have been spotted even in those with mild intermediate phenotypes—that is, in normal or relatively normal people who would fly under the radar of clinical significance for diagnosis of a personality disorder.39 One

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-related problem, all the time. The trouble is, it's difficult to conceive of an alternate, more refined shorthand that conveys the same sense of phenotype as Machiavellian. In point of fact, as with many personality types and disorders, there are almost certainly hundreds—perhaps thousands—of genotype configurations and resulting

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section, psychopaths, borderlines, Machiavellians, and the “successfully sinister” are often alluded to in virtually synonymous fashion. To some, this may seem an unfair blurring of phenotypes. To clarify matters, it might help if you were to think of psychopaths and borderlines as extreme examples, in slightly different but often overlapping fashion

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, of Machiavellian tendencies. The successfully sinister might be thought of as a less extreme form (“intermediate phenotype,” or subclinical manifestation) of these personality types. In a sense, then, all four of these terms blend into one another. b.Wine connoisseur Robert Parker

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and Minimum Body Mass Index,” Molecular Psychiatry 10, no. 9 (2005): 851–60. 29. J. Strauss et al., “BDNF and COMT Polymorphisms: Relation to Memory Phenotypes in Young Adults with Childhood-Onset Mood Disorder,” Neuromolecular Medicine 5, no. 3 (2004): 181–92. 30. A. Thapar et al., “Catechol O-methyltransferase Gene

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Attention-Deficit/Hyperactivity Disorder,” Archives of General Psychiatry 62, no. 11 (2005): 1275–78. 31. Ke Xu and D. Goldman, “Catechol-O-methyltransferase Genotype, Intermediate Phenotype, and Psychiatric Disorders,” in Cell Biology of Addiction, ed. Bertha Madras et al. (Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press, 2005), pp. 29

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,” Orthopedics 14, no. 11 (1991): 1185–93. 11. Ibid. CHAPTER 6: THE CONNECTION BETWEEN MACHIAVELLIANISM AND PERSONALITY DISORDERS 1. I. I. Gottesman, “Defining Genetically Informed Phenotypes for the DSM-V,” in Descriptions and Prescriptions: Values, Mental Disorders and the DSMs, ed. J. Z. Sadler (Baltimore, MD: Johns Hopkins University Press, 2002

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body's cells that provides the instructions to produce, for example, red hair. Also refers to the genetic makeup, as opposed to the physical appearance (phenotype), of a person. gray matter. The parts of neural tissue that are not covered with a white insulating substance called myelin. This contrasts with white

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, and function of organs such as the amygdala and cingulate cortex and then evaluating the same person's genes to see how they compare. intermediate phenotype. Phenotype generally refers to a trait you can see on the outside of a body, such as red hair. Intermediate

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phenotype is related to a trait or part on the inside of the body, such as the liver or the amygdala. Many times it's easier

…

to see if a gene has an effect on the intermediate phenotype inside the body than on the phenotype visible outside the body. junk DNA. Molecules of DNA that seem to serve no useful purpose and lie around generally being bored

…

in various parts of the brain. Although a purist would object, these images can be thought of as equivalent to color x-rays of tissues. phenotype. Phenotype generally refers to a trait you can see on the outside of a body, such as red hair. This contrasts with genotype, which is the

…

effect of immune system on, 207 evolutionary perspective on the disorder by psychiatrist Regina Pally, 282 general definition of, 135 heritability of, 85, 85n intermediate phenotype and, 66 litigation and unnecessary problems created by those with, 149 Medline's number of studies about, 33 neuroscience behind cognitive-perceptual impairment: dorsolateral, ventromedial

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Williams syndrome's abundance of, 98–99 emptiness, as DSM-IV diagnostic criteria for borderline personality disorder, 158 “end justifies means” behavior, 204 endophenotype. See phenotype, intermediate Engineering in Medicine and Biology Society, 211–12 English speakers versus Chinese speakers, neurological differences of, 175–76 Enron compared with sinister aspects of

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from, 60 fundamental overview of, 60–65 genetic drift, 65 “genetic stamping,” large effect of certain historical individuals, 267–70 “imaging genetics,” 67–69 intermediate phenotype and, 66–69 loss of olfactory and detoxification-related genes, 262, 262n of personality traits and disorders affective instability, heritability, 86 impulsivity, heritability, 86 lack

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traits), 295 Mao, 250 Shockley, William (also his disparaging comment about his children's), 290–91 Stalin, Joseph, 312 Stroessner, Alfredo (Paraguayan dictator), 311 intermediate phenotype. See also subclinical personality disorders cautions regarding this approach, 363n9 defined, 66–69, 255n in relation to “Cluster B” disorders and MAO-A gene, 238

…

, 217 results of his policies, 216–17, 248, 251–52 schizophrenia of son, 222–23, 227 MAO-A (monoamine oxidase A) alleles produce different intermediate phenotypes, 80 childhood stress and predisposition for antisocial behavior, 54, 80–82 Marcus Aurelius, 275, 275n Markovic, Mira (Slobodan Milosevic's wife and virtual Svengali), 155

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psychiatric effects of polio, 325–26 NIH (National Institutes of Health) overview of work mapping genetic bases of psychiatric illnesses, 68 research related to intermediate phenotype, 66–67 Nisbett, Richard E., codirector of University of Michigan's Culture and Cognition Program, 175 Nixon, Richard, temper of, 300 Niyazov, Saparmurat, dictator of

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, 197 Peterson, Laci, murdered by husband, previously dated violent man, 255 Phelps, Jim discussion of BDNF, 77–78 preface by, 19–22 phenotype explanation of, 64 intermediate. See intermediate phenotype Pinker, Steven: his The Blank Slate helped shift researchers away from “people are naturally good” idea, 37, 175 Pinochet, Augusto (president of

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, 64, 66–67 DARPP-32 and, 82–83 dopamine system and, 184 general discussion of, 227 gray matter loss with one type of, 205 intermediate phenotype and, 66–67 of Mao's son, 223, 227 placement in Axis I of DSM-IV, 134 theta rhythms and, 148 toxoplasma gondii and, 85

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

…

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

…

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

…

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

…

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

…

. 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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

, 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

…

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

…

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

…

, 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

…

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

…

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

…

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

…

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

…

, 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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

…

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

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