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MILL Iy Section impacts Genetics and general cognitive ability Robert Plomin General cognitive ability (g), often referred to as ‘general intelligence’, predicts social outcomes such as educational and occupational levels far better than any other behavioural trait. g is one of the most heritable behavioural traits, and genes that contribute to the heritability of g will certainly be identified. What are the scientific and social implications of finding genes associated with g? uring the past three decades, the behavioural sciences have emerged Dfrom an era of strict environmental explanations for differences in behaviour to a more balanced view that recognizes the importance of nature (genetics) as well as nurture (environment). This shift occurred first for behavioural disorders, including rare disorders such as autism (which has an incidence of 1 per 1,000 population), more common disorders such as schizophrenia (1 in 100), and very common disorders such as reading disability (1 in 50). More recently, it has become increasingly accepted that genetic variation makes an important con- tribution to differences among individuals in the normal range of behaviour as well as for abnormal behaviour. Moreover, many behavioural disorders, especially common ones, may represent variation at the extremes of the same genetic and environ- mental factors that are responsible for varia- Figure 1 Quantitative trait locus (QTL) perspective on complex traits. Differences among individuals tion within the normal range. For example, for most quantitative or complex traits such as reading ability are distributed as a normal bell-shaped disorders such as reading disability may not curve. Multiple genes influence complex traits as probabilistic propensities rather than be due to genetic variants that specifically predetermined programmes. Here the different genetic make-up of individuals with respect to two influence the disorder. Rather, the hypothetical genes involved in reading ability is shown for 100 individuals (each person is same genes that contribute to the normal represented by an oval), with five of these individuals (those on the extreme left) receiving a diagnosis range of individual differences in reading of reading disability. The green ovals indicate that the individual has the disabling variant of one gene ability may be responsible for reading and blue ovals denote the disabling variant of the other gene. Neither gene is necessary or sufficient disability (Fig. 1). This view, known as the for low scores, even for individuals who have disabling variants of both genes (red ovals). This QTL quantitative trait locus (QTL) perspective perspective suggests that genes associated with common disorders such as reading disability may (see Box 1), has important implications for represent the quantitative extreme of the same genes that are responsible for variation throughout the search for the genes responsible for the population. behaviour because such genes will individu- ally have small effects; this will make them more difficult to find than genes that have memory — correlate substantially with each lay people often read in the popular press major effects1. other, and general cognitive ability (g) is that the assessment of intelligence is what these diverse measures have in circular — intelligence is what intelligence General cognitive ability common (see Box 2). Clearly there is more to tests assess. On the contrary, g is one of the For historical and political reasons, one cognition than g — although g explains most reliable and valid measures in the quantitative trait in particular is highly about 40 per cent of the variance among such behavioural domain; its long-term stability controversial. This is general cognitive tests, most of the variance of a particular test after childhood is greater than for any other ability, which has a normal distribution in is independent of g. behavioural trait, and it predicts important the population from a low end of mild There is a wide gap between what lay social outcomes such as educational and mental handicap to a high end of gifted indi- people (including scientists in other occupational levels far better than any other viduals. Diverse measures of cognitive fields) believe about intelligence and intelli- trait2. Although a few critics remain3, g is abilities — such as spatial ability, verbal gence testing, and what the professional widely accepted by experts4. But it is less clear ability, information processing speed and behavioural scientist believes. Most notably, what g is: is it due to a single general process NATURE | VOL 402 | SUPP | 2 DECEMBER 1999 | www.nature.com C25 impacts ty’, vary from 40 to 80 per cent, but estimates family members for gare important in child- based on the entire body of data make it hood, accounting for about a quarter of about 50 per cent, indicating that genes the variance, but they are not important account for about half of the variance in g. after adolescence. In other words, non- When the data are sorted by age, heritability shared environmental factors that make is found to increase from about 20 per cent in children in the same family different (such infancy, to about 40 per cent in childhood, to as differences in parental treatment, differ- 60 per cent or greater later in life12, even for ences in school experiences and different 13 individuals over 80 years old . This increase experiences with peers) provide the long- in heritability throughout lifespan is inter- term consequences of environmental esting, because it is counterintuitive to the influence for g. This finding coincides with effects of Shakespeare’s ‘slings and arrows of similar findings for other quantitative traits outrageous fortune’ accumulating over and indicates the need to re-examine the 16 time. It may be that heritability increases nurture assumption . because individuals seek out and create Two other examples of recent genetic environments correlated with their genetic findings about the environment are that propensities. environmental influences may override Most of the genetic variance for g is addi- genetic effects in families of low socioeco- 17 tive, that is, the effects of the individual genes nomic status , and that genetic factors seem simply to add up rather than there contribute to individuals’ interactions with being interactions between the genes. The their environment18. The former finding additivity of most genetic effects on g may be highlights that heritability estimates are not because there is greater assortative mating absolute but depend on the environment (non-random mating) for g than for any in which they are measured. The latter other behavioural trait. In other words, observation, called ‘genotype–environment bright women are likely to mate with bright correlation’, indicates that genetic influences Figure 2 Functional genomics includes all levels men and the outcome of this dual effect is on abilities can best be thought of as of analysis from molecular biology to that their offspring are likely to be brighter psychology. The higher levels of analysis can be on average than would be expected if mating referred to as behavioural genomics in order to were at random, thus spreading out the emphasize the importance of top-down analyses distribution of g in the population. Box 1The source of of pathways between genes and behaviour. The data that provide evidence for a genetic effect on g also provide the best genetic variation available evidence for the importance of such as high-level strategies called executive environmental factors that are independent Quantitative traits are those characteristics, such function or speed of information processing, of genetics. Environment clearly is impor- as height or ‘intelligence’, that are found as a or does it represent a concatenation of more tant, as indicated by the steady rise in IQ continuum of values within a population rather specific cognitive processes?5,6 scores during the past several generations, than as the discrete alternative inherited character The concept of a genetic contribution to g which would seem too short a time to be states familiar to most people from a schoolroom has provoked much controversy, especially explained by genetics14, and by studies in acquaintance with Mendel’s peas. They are due to following the publication in 1994 of The Bell which children from abusive families show the combined effects of a number of different Curve by Herrnstein and Murray7. (This gains in IQ when adopted15. genes (each of which will, of course, be inherited book in fact scarcely touches on genetics and according to the rules of mendelian genetics), and does not view genetic evidence as crucial to Recent findings and new directions usually also have a considerable environmental its arguments.) The first half of the book Genetic research has moved beyond the input to the final outcome. This final outcome is shows, like many other studies, that g is rudimentary questions of whether, and to known as the phenotype. A quantitative trait locus related to educational and social outcomes, what extent, genetic differences are impor- (QTL) refers to a gene that contributes to a but the second half attempts to argue that tant in the origins of individual differences in quantitative trait. A locus is the technical name in certain right-wing policies follow from these g. These new findings inform the scientific genetics for the position on the chromosome at findings. However, as discussed later in this and social implications
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