Heritability of IQ

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Study of the heritability of IQ is a controversial field of research that includes biology, psychology, philosophy, sociology, and anthropology. Heritability is a measure of the relative contribution of genes to the variation of a phenotype on a given group in a specific environment.^[1] The debate about IQ heritability touches on the nature versus nurture divide,^[2] and there has been no consensus in the academic community about it even since research began in the 19th century.^[3] Most of the heritable variance in IQ appears to be carried by the general intelligence factor (or g). IQ is a polygenic trait under normal circumstances according to recent research.^[4] However, destructive mutation of individual genes associated with development can severely affect intelligence, with Phenylketonuria as an example.^[5]

Throughout the developed world, specific estimates in the academic research into the heritability of IQ have varied from relatively high, such as over 0.9 in a 1994 report,^[6] to relatively low, such as below 0.5 in a 1997 report.^[3] A 1996 statement by the American Psychological Association gave about .45 for children and about .75 during and after adolescence.^[7] A 2004 meta-analysis of reports in Current Directions in Psychological Science gave an overall estimate of around three quarters.^[8] The New York Times Magazine has also listed about three quarters as a figure held by the majority of studies.^[9] This coefficient would imply that r squared is about 0.56, meaning that about 56% of the variance in IQ scores is genetic.

Contents 1 Methods and results 1.1 Heritability calculations 1.1.1 Background 1.1.2 Estimates and caveats to them 1.2 Developing nations 2 Issues in the calcuations 2.1 Family environment 2.2 Biased older studies? 2.3 Maternal (fetal) environment 2.4 The Dickens and Flynn model 2.5 Regression towards the mean 2.6 Historical research 2.7 The inheritance of cognitive deficits 3 The search for specific genes 4 Between-group heritability 5 Literature 6 References

[edit] Methods and results

[edit] Heritability calculations

[edit] Background

Heritability is defined as the proportion of variance in a trait which is attributable to genes within a defined population in a specific environment.^[1] Heritability takes a value ranging from 0 to 1; a heritability of 1 indicates that all variation in the trait in question is genetic in origin and a heritability of 0 indicates that none of the variation is genetic. The heritability of many traits can be considered primarily genetic under similar environmental backgrounds. For example, Visscher et al. (2006) found that adult height has a heritability estimated at 0.80, when a relatively uniform environmental background is present, to control for environment the study only looked at the contribution of heritability to variation within families. The paper stated that "one can never be sure that the estimates are correct, because nature and nurture can be confounded without one knowing it. The authors got around this problem by comparing the similarity between relatives as a function of the exact proportion of genes that they have in common, looking only within families."^[2] Other traits have low heritabilities, which indicate a large relative environmental influence. For example, a twin study on the heritability of depression in men calculated it as 0.29, while it was 0.42 for women in the same study.^[10]

Heritability for a trait is calculated by measuring how strongly traits covary in people of a given genetic and environmental similarity. The most common method is to consider identical twins reared apart, with any difference which exists between such twin pairs only attributed to the environment. In terms of correlation statistics, this means that theoretically the correlation of tests scores between monozygotic twins would be 1.00 if genetics alone accounted for variation in IQ scores; likewise, siblings and dizygotic twins share half of their alleles and the correlation of their scores would be 0.50 if IQ were affected by genes alone. Practically, however, the upper bound of these correlations are given by the reliability of the test, which tends to be 0.90 to 0.95 for typical IQ tests^[11] Thus, the actual heritability of IQ will tend to be slightly higher than attained by estimates derived from studies of monozygotic twins, though this effect is small.

In the case of the inheritance of IQ or a certain degree of giftedness, the relatives of probands with a high IQ exhibit a comparably high IQ with a much higher probability than the general population. In 1982, Bouchard and McGue reviewed such correlations reported in 111 original studies in the United States.^[12] The mean correlation of IQ scores between monozygotic twins was 0.86, between siblings, 0.47, between half-siblings, 0.31, and between cousins, 0.15. From such data the heritability of IQ was estimated at anywhere between 0.40 and 0.80 in the United States. The reason for this wide margin appeared to be that the heritability of IQ rises through childhood and adolescence, peaking at 0.68 and 0.78 in adults, leaving the overwhelming majority of IQ differences between individuals to be explained genetically.^[13]

The finding of rising heritability with age is counter-intuitive; it is reasonable to expect that genetic influences on traits like IQ should become less important as one gains experiences with age. However, that the opposite occurs is well documented. According to work by Robert Plomin,heritability estimates calculated on infant samples are as low as 20%, rising to around 40% in middle childhood, and ultimately as high as 80% in adult samples in the United States.^[14] This suggests that the underlying genes actually express themselves by affecting a person's predisposition to build, learn, and develop mental abilities throughout the lifespan.^{[citation needed]}

[edit] Estimates and caveats to them

In 2006, The New York Times Magazine listed about three quarters as a figure held by the majority of studies.^[9] A 2004 meta-analysis of reports in Current Directions in Psychological Science gave an estimate of around three quarters as well.^[8] As well, a 1996 statement by the American Psychological Association gave about .45 for children and about .75 during and after adolescence.^[7] The 2005 edition of Assessing adolescent and adult intelligence by Alan S. Kaufman and Elizabeth O. Lichtenberger found correlations of 0.86 for identical twins raised together compared to 0.76 for those raised apart and 0.47 for siblings.^[15] A 1994 review in Behavior Genetics based on identical/fraternal twin studies found that it is as high as 0.92 in general cognitive ability but it also varies based on the trait, with .60 for verbal tests, .50 for spatial and speed-of-processing tests, and only .40 for memory tests.^[6]

There are a number of points to consider when interpreting heritability:

• A high heritability does not mean that the environment has no effect on the development of a trait, or that learning is not involved. Vocabulary size, for example, is very substantially heritable (and highly correlated with general intelligence) although every word in an individual's vocabulary is learned. In a society in which plenty of words are available in everyone's environment, especially for individuals who are motivated to seek them out, the number of words that individuals actually learn depends to a considerable extent on their genetic predispositions.^[7]
• A common error is to assume that because something is heritable it is necessarily unchangeable. This is wrong. Heritability does not imply immutability. As previously noted, heritable traits can depend on learning, and they may be subject to other environmental effects as well. The value of heritability can change if the distribution of environments (or genes) in the population is substantially altered. For example, an impoverished or suppressive environment could fail to support the development of a trait, and hence restrict individual variation. This could affect estimates of heritability.^[7] Another example is Phenylketonuria which previously caused mental retardation for everyone who had this genetic disorder. Today, this can be prevented by following a modified diet.
• On the other hand, there can be effective environmental changes that do not change heritability at all. If the environment relevant to a given trait improves in a way that affects all members of the population equally, the mean value of the trait will rise without any change in its heritability (because the differences among individuals in the population will stay the same). This has evidently happened for height: the heritability of stature is high, but average heights continue to increase.^[7]
• Even in developed nations, high heritability of a trait within a given group has no necessary implications for the source of a difference between groups.^[7][16]
• Heritability increases with education and with social class, such that the children of highly educated parents are more likely to inherent traits.^[15][9] Most of the studies done about heritability have used middle class families- not because of any intentional choice made by researchers but because, on average, lower income people are less likely to volunteer to be studied in the first palce.^[9]

[edit] Developing nations

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See also: Health and intelligence

Almost all studies on heritability have been in the developed world, mostly in the United States.^{[citation needed]} In developing nations there are many environmental factors affecting IQ which are much less important in developed nations.^{[citation needed]} Examples include nutrition, diseases, environmental toxins, and health care.^{[citation needed]} This likely affects heritability.^{[citation needed]}

[edit] Issues in the calcuations

[edit] Family environment

In the developed world, nearly all personality traits show that, contrary to expectations, environmental effects actually cause non-related children raised in the same family ("adoptive siblings") to be as different as children raised in different families (Harris, 1998; Plomin & Daniels, 1987). There are some family effects on the IQ of children, accounting for up to a quarter of the variance. However, by adulthood, this correlation disappears, such that adoptive siblings are not more similar in IQ than strangers,^[17] while adult full siblings show an IQ correlation of 0.6. Twin studies reinforce this pattern: monozygotic (identical) twins raised separately are highly similar in IQ (0.86), more so than dizygotic (fraternal) twins raised together (0.6) and much more than adoptive siblings (~0.0).^[18]

The American Psychological Association's report Intelligence: Knowns and Unknowns (1995) states that there is no doubt that normal child development requires a certain minimum level of responsible care. Severely deprived, neglectful, or abusive environments must have negative effects on a great many aspects of development, including intellectual aspects. Beyond that minimum, however, the role of family experience is in serious dispute. Do differences between children's family environments (within the normal range) produce differences in their intelligence test performance? The problem here is to disentangle causation from correlation. There is no doubt that such variables as resources of the home and parents' use of language are correlated with children's IQ scores, but such correlations may be mediated by genetic as well as (or instead of) environmental factors. But how much of that variance in IQ results from differences between families, as contrasted with the varying experiences of different children in the same family? Recent twin and adoption studies suggest that while the effect of the family environment is substantial in early childhood, it becomes quite small by late adolescence. These findings suggest that differences in the life styles of families whatever their importance may be for many aspects of children's lives make little long-term difference for the skills measured by intelligence tests. It also stated "We should note, however, that low-income and non-white families are poorly represented in existing adoption studies as well as in most twin samples. Thus it is not yet clear whether these studies apply to the population as a whole. It remains possible that, across the full range of income and ethnicity, between-family differences have more lasting consequences for psychometric intelligence."^[19]

A study of French children adopted between the ages of 4 and 6 shows the continuing interplay of nature and nurture. The children came from poor backgrounds with IQs that initially averaged 77, putting them near retardation. Nine years later after adoption, they retook the IQ tests, and all of them did better. The amount they improved was directly related to the adopting family’s status. "Children adopted by farmers and laborers had average IQ scores of 85.5; those placed with middle-class families had average scores of 92. The average IQ scores of youngsters placed in well-to-do homes climbed more than 20 points, to 98."^[9]

[edit] Biased older studies?

Stoolmiller (1999) found that the range restriction of family environments that goes with adoption, that adopting families tend to be more similar on for example SES than the general population, means that role of the shared family environment have been underestimated in previous studies. Corrections for range correction applied to adoption studies indicate that SE could account for as much as 50% of the variance in IQ.^[20] However, the effect of restriction of range on IQ for adoption studies was examined by Matt McGue and colleagues, who write that "restriction in range in parent disinhibitory psychopathology and family SES had no effect on adoptive-sibling correlations [in] IQ".^[21]

Eric Turkheimer and colleagues (2003), not using an adoption study, included impoverished US families. Results demonstrated that the proportions of IQ variance attributable to genes and environment vary nonlinearly with SES. The models suggest that in impoverished families, 60% of the variance in IQ is accounted for by the shared family environment, and the contribution of genes is close to zero; in affluent families, the result is almost exactly the reverse.^[22] They suggest that the role of shared environmental factors may have been underestimated in older studies which often only studied affluent middle class families.^[23]

When comparing late 1970s to pre-1963 recorded data, researches DeFries and Plomin found that IQ correlation between parent and child living together fell significantly, from 0.50 to 0.35.^[15]

[edit] Maternal (fetal) environment

A meta-analysis, by Devlin and colleagues in Nature (1997), of 212 previous studies evaluated an alternative model for environmental influence and found that it fits the data better than the 'family-environments' model commonly used. The shared maternal (foetal) environment effects, often assumed to be negligible, account for 20% of covariance between twins and 5% between siblings, and the effects of genes are correspondingly reduced, with two measures of heritability being less than 50%. They argue that the shared maternal environment may explain the striking correlation between the IQs of twins, especially those of adult twins that were reared apart.^[3]

Bouchard and McGue reviewed the literature in 2003, arguing that Devlin's conclusions about the magnitude of heritability is not substantially different than previous reports and that their conclusions regarding prenatal effects stands in contradiction to many previous reports.^[24] They write that:

Chipuer et al. and Loehlin conclude that the postnatal rather than the prenatal environment is most important. The Devlin et al. (1997a) conclusion that the prenatal environment contributes to twin IQ similarity is especially remarkable given the existence of an extensive empirical literature on prenatal effects. Price (1950), in a comprehensive review published over 50 years ago, argued that almost all MZ twin prenatal effects produced differences rather than similarities. As of 1950 the literature on the topic was so large that the entire bibliography was not published. It was finally published in 1978 with an additional 260 references. At that time Price reiterated his earlier conclusion (Price, 1978). Research subsequent to the 1978 review largely reinforces Price’s hypothesis (Bryan, 1993; Macdonald et al., 1993; Hall and Lopez-Rangel, 1996; see also Martin et al., 1997, box 2; Machin, 1996).^[25]

[edit] The Dickens and Flynn model

Dickens and Flynn (2001) argue that the arguments regarding the disappearance of the shared family environment should apply equally well to groups separated in time. This is contradicted by the Flynn effect. Changes here have happened too quickly to be explained by genetics. This paradox can be explained by observing that the measure "heritability" includes both a direct effect of the genotype on IQ and also indirect effects where the genotype changes the environment, in turn effecting IQ. That is, those with a higher IQ tend to seek out stimulating environments that further increase IQ. The direct effect can initially have been very small but feedback loops can create large differences in IQ. In their model an environmental stimulus can have a very large effect on IQ, even in adults, but this effect also decays over time unless the stimulus continues (the model could be adapted to include possible factors, like nutrition in early childhood, that may cause permanent effects). The Flynn effect can be explained by a generally more stimulating environment for all people. The authors suggest that programs aiming to increase IQ would be most likely to produce long-term IQ gains if they taught children how to replicate outside the program the kinds of cognitively demanding experiences that produce IQ gains while they are in the program and motivate them to persist in that replication long after they have left the program.^[26][27]

[edit] Regression towards the mean

The heritability of IQ measures the extent to which the IQ of a child is measurably influenced by the IQ its parents. As IQ is a quantifiable phenotype, one can estimate the expected IQ of child using the equation , where

• is the expected IQ of the child,
• is the mean IQ of the population to which the parents belong,
• h² is the heritability of IQ,
• m and f are the IQs of the mother and father, respectively.^[28]

The equation asserts that, on average, the IQ of a child tends to the mean IQ of the population. For instance, if the heritability of IQ is 50% and the mean IQ of a population is 100, then a couple with an average IQ of 120 will, on average, have a child with an IQ of 110. Similarly, a couple with an average IQ of 80 will, on average, have a child with an IQ of 90.

It is noted that the above equation relates only statistical averages and is not deterministic. Furthermore, the equation is a general equation based in the inheritance of genetically-based characteristics (in this case, phenotypes), and so it is implicitly assumed that environmental factors are, for the sake of correctly assessing the genetic contribution to IQ, the same across the population.

Operating under the assumption that child and parent are raised in the exact same environment (unlikely, but usually closer to the truth than in the completely dissimilar environment that the previous equation assumes), h² can be replaced by h, which is simply the correlation between parent and offspring IQ. In this case, regression towards the mean is no longer partially caused by environmental differences and therefore only by random genetic variation.

Finally, it is important to note that the expected IQ of the offspring is normally distributed around the mean calculated using the above equation, so in many cases regression towards the mean does not actually occur; as the values are normally distributed, there is a chance that offspring IQ will be more deviant from the mean than that of the parental average.

[edit] Historical research

As early as 1869, Francis Galton replaced mere speculations by statistical data through his book, Hereditary Genius:

---

Highly Gifted Men and the Percentage of their Highly Gifted Male Relatives

(classified by occupation and achievement)

	Galton	Terman	Brimhall	Weiss
	%	%	%	%	n (Weiss)
Probands	100	84+	100	97+	1972: 1329 1994:   357
Fathers	26	41	29	40	346
Brothers	47	-	49	49	220
Sons	60	64*	-	55	77
Grandfathers	14	-	9	9	681
Uncles	16	-	13	14	615
Nephews	23	-	-	22	76
Grandsons	14	-	-	-	-
Greatgrandfathers	0	-	-	4	1290
Uncles of the parents	5	-	-	5	1996
Cousins	16	-	9#	18	570
Greatgrandsons	7	-	-	-	-
Cousins of parents	-	-	-	11	2250
"+": classified by occupation; 100%, if classified by test "*": classified only by IQ; classification by occupation gives about 55%; n = 820. "#": some cousins were still too young and did not have full opportunity to become distinguished "-": no data Sources:''' Francis Galton: Hereditary Genius. London 1869, p. 195.[1]. 100 famous Famous men (n = 43) of science and the percentage of their famous male relatives. M. H. Oden: The fulfillment of promise: 40-year follow-up of the Terman gifted group. Genetical Psychology Monographs 77 (1968) 3-93. The mean IQ (transformed to 100;15) of the sample of probands was 146 (n = 724); the cut-off score IQ 137. Dean R. Brimhall: Family resemblances among American men of science. The American Naturalist 56 (1922) 504-547; 57 (1923) 74-88, 137-152, and 326-344. In 1915 questionnaires were filled in by 956 distinguished American men of science and their relatives. Volkmar Weiss: Mathematical giftedness and family relationship. European Journal for High Ability 5 (1994) 58-67.[2] Highly gifted males (mean IQ 135 +/- 9) and their relatives in professions and occupational positions, typically associated with an IQ above 123.

Despite the differences in methods and societies, there is a notable parallelism in the published statistics. The ITO-method by Li and Sacks (1954) allows from this set of data the estimation of the underlying number of genes and their allele frequencies.

[edit] The inheritance of cognitive deficits

There are many genetic variants known to cause lower IQ. The number of such mutations already known is in the hundreds. For example, an allele of the gene GDI1 is associated with an IQ below 70.^{[citation needed]}

Copy number variation has also been associated with idiopathic learning disability. ^[29]

There are number of known cases where the homozygotes have severe cognitive deficits and the heterozygotes show a small decrease of IQ. In such cases further alleles are investigated to estimate their influence on IQ. For example, one minor allele of the gene ALDH5A1 is associated with an IQ difference of around 1.5 points.^[30]

Interindividual (between individuals) differences in learning ability are also known in mice, dogs and other animals, and the achievements of pure strains can be improved by selective breeding.^{[clarification needed]} In such a way also behaviour genetics is contributing to our knowledge of the inheritance of mental traits. There is an open question to which degree differences of animal behaviour have any meaning for differences in human intelligence.^{[clarification needed]}

[edit] The search for specific genes

Unfortunately, most of the research done about the heritability of intelligence have focused on children and young adults. Thus, the role of genetic factors to intelligence, which most likely increases over time, is mostly unknown.^[6] Many studies attempting to find loci in the genome relating to IQ have had little success. For example, a study by Robert Plomin using groups of around 100 people investigated 1,842 DNA markers in a high-IQ group and in an average-IQ control group. The study used a five-step replication process to eliminate false positives, and no gene met this rigid criterion for replicability^[31]

The failure to find a specific gene associated with IQ indicates that cognitive abilities are very complex and are likely to involve several genes (polygenic). Some estimate that as much as 40% of all genes may contribute to IQ.^[32] The more genes that contribute to a trait the more the trait will be continuous instead of discrete. A 2008 study of 500,000 single nucleotide polymorphisms (SNPs) from 7,089 children did not substantially improve on earlier studies. The study did not find any any SNPs that accounted for more than 0.5% of the variance in general intelligence.^[33]

A recent study did find that a gene called FADS2 along with breastfeeding adds about 7 IQ points to those with the "C" version of the gene. Those with the "G" version see no advantage.^[34][35]

There is "a highly significant association" between the CHRM2 gene and intelligence according to a 2006 Dutch family study. The study concluded that there was an association between the CHRM2 gene on chromosome 7 and Performance IQ, as measured by the Wechsler Adult Intelligence Scale-Revised. The Dutch family study used a sample of 667 individuals from 304 families.^[36] A similar association was found independently in the Minnesota Twin and Family Study (Comings et al. 2003) and by the Department of Psychiatry at the Washington University.^[37] Microcephalin and ASPM are two genes that are associated with brain development. Mutations in these genes are associated with microcephaly, and hence they were initially associated with general intelligence. However recent studies have found no association with general cognitive abilities.^[38]

[edit] Between-group heritability

The fact that IQ differences between individuals are found to have a very large genetic component does not necessitate that mean group-level disparities in IQ must likewise have a genetic basis. An analogy, attributed to Richard Lewontin,^[39] illustrates this point:

Suppose two handfuls are taken from a sack containing a genetically diverse variety of corn, and each grown under carefully controlled and standardized conditions, except that one batch is lacking in certain nutrients that are supplied to the other. After several weeks, the plants are measured. There is variability of growth within each batch, due to the genetic variability of the corn. Given that the growing conditions are closely controlled, nearly all the variation in the height of the plants within a batch will be due to differences in their genes. Thus, within populations, heritabilities will be very high. Nevertheless, the difference between the two groups is due entirely to an environmental factor - differential nutrition. Lewontin didn't go so far as to have the one set of pots painted white and the other set black, but you get the idea. The point of the example, in any case, is that the causes of between-group differences may in principle be quite different from the causes of within-group variation.^[40]

In response to Lewontin, Arthur Jensen maintained that environmental differences are too small to account for between-group IQ differences, and that therefore genetic differences must provide the primary explanation.^[41]

[edit] Literature

• McGuffin P (2000). "The quantitative and molecular genetics of human intelligence". Novartis Found. Symp. 233: 243–55; discussion 255–9, 276–80. doi:10.1002/0470870850.ch15. PMID 11276906. 

• G. Meisenberg: Genes for intelligence. A review of recent progress. Mankind Quarterly 36 (Winter 2005) 139-164. abstract

• Payton A (2006). "Investigating cognitive genetics and its implications for the treatment of cognitive deficit". Genes, Brain and Behavior 5 Suppl 1: 44–53. doi:10.1111/j.1601-183X.2006.00194.x. PMID 16417617. 

[edit] References

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