122:
variation decreases, causing individuals to show less phenotypic variation, like showing more similar levels of intelligence. Heritability increases when genetics are contributing more variation or because non-genetic factors are contributing less variation; what matters is the relative contribution. Heritability is specific to a particular population in a particular environment. High heritability of a trait, consequently, does not necessarily mean that the trait is not very susceptible to environmental influences. Heritability can also change as a result of changes in the environment, migration,
4105:
27:
5155:
117:. This is not the same as saying that this fraction of an individual phenotype is caused by genetics. For example, it is incorrect to say that since the heritability of personality traits is about 0.6, that means that 60% of your personality is inherited from your parents and 40% comes from the environment. In addition, heritability can change without any genetic change occurring, such as when the environment starts contributing to more variation. As a case in point, consider that both
514:
105:). It is the source of much confusion due to the fact that its technical definition is different from its commonly-understood folk definition. Therefore, its use conveys the incorrect impression that behavioral traits are "inherited" or specifically passed down through the genes. Behavioral geneticists also conduct heritability analyses based on the assumption that genes and environments contribute in a separate, additive manner to behavioral traits.
5462:...all complex human traits result from a combination of causes. If these causes interact, it is impossible to assign quantitative values to the fraction of a trait due to each, just as we cannot say how much of the area of a rectangle is due, separately, to each of its two dimensions. Thus, in the analyses of complex human phenotypes...we cannot actually find 'the relative importance of genes and environment in the determination of phenotype'.
3598:
3888:. Eric Turkheimer has argued that newer molecular methods have vindicated the conventional interpretation of twin studies, although it remains mostly unclear how to explain the relations between genes and behaviors. According to Turkheimer, both genes and environment are heritable, genetic contribution varies by environment, and a focus on heritability distracts from other important factors. Overall, however,
3783:
830:
1644:
3849:, and that this alleged bias distracts from other factors that researches have found more causally important, such as childhood abuse causing later psychosis. Heritability estimates are also inherently limited because they do not convey any information regarding whether genes or environment play a larger role in the development of the trait under study. For this reason,
493:. If a selective pressure such as improving livestock is exerted, the response of the trait is directly related to narrow-sense heritability. The mean of the trait will increase in the next generation as a function of how much the mean of the selected parents differs from the mean of the population from which the selected parents were chosen. The observed
3503:
individuals across a broad range of environments, although inference of genetic variance from phenotypic and environmental variance may lead to underestimation of heritability due to the challenge of capturing the full range of environmental influence affecting a trait. Other methods for calculating heritability use data from
579:
3584:
When genome-wide genotype data and phenotypes from large population samples are available, one can estimate the relationships between individuals based on their genotypes and use a linear mixed model to estimate the variance explained by the genetic markers. This gives a genomic heritability estimate
1524:
who have been separated early in life and raised in different environments. Such individuals have identical genotypes and can be used to separate the effects of genotype and environment. A limit of this design is the common prenatal environment and the relatively low numbers of twins reared apart. A
385:
to each offspring, parent-offspring resemblance depends upon the average effect of single alleles. Additive variance represents, therefore, the genetic component of variance responsible for parent-offspring resemblance. The additive genetic portion of the phenotypic variance is known as Narrow-sense
164:
A prerequisite for heritability analyses is that there is some population variation to account for. This last point highlights the fact that heritability cannot take into account the effect of factors which are invariant in the population. Factors may be invariant if they are absent and do not exist
3696:
For example, imagine that a plant breeder is involved in a selective breeding project with the aim of increasing the number of kernels per ear of corn. For the sake of argument, let us assume that the average ear of corn in the parent generation has 100 kernels. Let us also assume that the selected
1655:
Heritability may be estimated by comparing parent and offspring traits (as in Fig. 2). The slope of the line (0.57) approximates the heritability of the trait when offspring values are regressed against the average trait in the parents. If only one parent's value is used then heritability is twice
1707:
Heritability for traits in humans is most frequently estimated by comparing resemblances between twins. "The advantage of twin studies, is that the total variance can be split up into genetic, shared or common environmental, and unique environmental components, enabling an accurate estimation of
1685:– the sum of half the additive genetic variance plus full effect of the common environment. It thus places an upper limit on additive heritability of twice the full-Sib phenotypic correlation. Half-Sib designs compare phenotypic traits of siblings that share one parent with other sibling groups.
121:
and environment have the potential to influence intelligence. Heritability could increase if genetic variation increases, causing individuals to show more phenotypic variation, like showing different levels of intelligence. On the other hand, heritability might also increase if the environmental
3857:
describe the term "heritability" in the context of behavior genetics as "...one of the most misleading in the history of science" and argue that it has no value except in very rare cases. When studying complex human traits, it is impossible to use heritability analysis to determine the relative
3530:
The currently popular methodology relies on high degrees of certainty over the identities of the sire and dam; it is not common to treat the sire identity probabilistically. This is not usually a problem, since the methodology is rarely applied to wild populations (although it has been used for
88:
Heritability is estimated by comparing individual phenotypic variation among related individuals in a population, by examining the association between individual phenotype and genotype data, or even by modeling summary-level data from genome-wide association studies (GWAS). Heritability is an
3502:
A wide variety of approaches using linear mixed models have been reported in literature. Via these methods, phenotypic variance is partitioned into genetic, environmental and experimental design variances to estimate heritability. Environmental variance can be explicitly modeled by studying
3692:
In this equation, the
Response to Selection (R) is defined as the realized average difference between the parent generation and the next generation, and the Selection Differential (S) is defined as the average difference between the parent generation and the selected parents.
2132:
3697:
parents produce corn with an average of 120 kernels per ear. If h equals 0.5, then the next generation will produce corn with an average of 0.5(120-100) = 10 additional kernels per ear. Therefore, the total number of kernels per ear of corn will equal, on average, 110.
85:. In human studies of heritability these are often apportioned into factors from "shared environment" and "non-shared environment" based on whether they tend to result in persons brought up in the same household being more or less similar to persons who were not.
1393:" unaccounted for by known genetic loci, the assumption of additivity may render these estimates invalid. There is also some empirical evidence that the additivity assumption is frequently violated in behavior genetic studies of adolescent intelligence and
825:{\displaystyle {\begin{aligned}P_{ij}&=\mu +\alpha \,(B_{i}+B_{j})+\delta \,(B_{i}B_{j})\\&={\text{Population mean}}+{\text{Additive Effect }}(a_{ij}=\alpha (B_{i}+B_{j}))+{\text{Dominance Deviation }}(d_{ij}=\delta (B_{i}B_{j})).\\\end{aligned}}}
1700:
1730:, contributes to similarity between siblings due to the commonality of the environment they are raised in. Shared environment is approximated by the DZ correlation minus half heritability, which is the degree to which DZ twins share the same genes,
1680:
A basic approach to heritability can be taken using full-Sib designs: comparing similarity between siblings who share both a biological mother and a father. When there is only additive gene action, this sibling phenotypic correlation is an index of
3511:) on the trait. This can lead to underestimation of heritability, however. This discrepancy is referred to as "missing heritability" and reflects the challenge of accurately modeling both genetic and environmental variance in heritability models.
1432:
In non-human populations it is often possible to collect information in a controlled way. For example, among farm animals it is easy to arrange for a bull to produce offspring from a large number of cows and to control environments. Such
481:
For traits which are not continuous but dichotomous such as an additional toe or certain diseases, the contribution of the various alleles can be considered to be a sum, which past a threshold, manifests itself as the trait, giving the
3526:
will usually have three or more data points for each individual: a code for the sire, a code for the dam and one or several trait values. Different trait values may be for different traits or for different time points of measurement.
126:, or the way in which heritability itself is measured in the population under study. The heritability of a trait should not be interpreted as a measure of the extent to which said trait is genetically determined in an individual.
3589:. Particularly, the method called High-Definition Likelihood (HDL) can estimate genomic heritability using only GWAS summary statistics, making it easier to incorporate large sample size available in various GWAS meta-analysis.
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987:
1388:
Estimates of the total heritability of human traits assume the absence of epistasis, which has been called the "assumption of additivity". Although some researchers have cited such estimates in support of the existence of
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148:
to help to identify the causes of differences between individuals. Since heritability is concerned with variance, it is necessarily an account of the differences between individuals in a population. Heritability can be
153:– examining a single trait – or multivariate – examining the genetic and environmental associations between multiple traits at once. This allows a test of the genetic overlap between different phenotypes: for instance
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465:
350:
69:
between individuals in that population. The concept of heritability can be expressed in the form of the following question: "What is the proportion of the variation in a given trait within a population that is
2395:
5365:
2785:
3531:
several wild ungulate and bird populations), and sires are invariably known with a very high degree of certainty in breeding programmes. There are also algorithms that account for uncertain paternity.
1884:
Consider an experiment with a group of sires and their progeny from random dams. Since the progeny get half of their genes from the father and half from their (random) mother, the progeny equation is
1712:
for the trait), and so identical or monozygotic (MZ) twins on average are twice as genetically similar as DZ twins. A crude estimate of heritability, then, is approximately twice the difference in
1372:
1103:
3763:
1614:
584:
1982:
1947:
5643:
1421:
components of variance depend on the sample characteristics. Briefly, better estimates are obtained using data from individuals with widely varying levels of genetic relationship - such as
2678:. The expected mean square is calculated from the relationship of the individuals (progeny within a sire are all half-sibs, for example), and an understanding of intraclass correlations.
2508:
133:
a phenotype, making its expression almost inevitable in all occurring environments. Individuals with the same genotype can also exhibit different phenotypes through a mechanism called
1544:. Depending on the methods used to estimate heritability, correlations between genetic factors and shared or non-shared environments may or may not be confounded with heritability.
3011:
556:
are either 0 or 1, the expected phenotype can then be written as the sum of the overall mean, a linear effect, and a dominance deviation (one can think of the dominance term as an
1409:
can be observed or measured directly, heritability must be estimated from the similarities observed in subjects varying in their level of genetic or environmental similarity. The
2550:
1822:
5928:
3490:
3462:
3408:
3375:
3347:
3293:
3239:
4133:
3845:. Bentall has claimed that such heritability scores are typically calculated counterintuitively to derive numerically high scores, that heritability is misinterpreted as
1762:
We use the basic discussion of
Kempthorne. Considering only the most basic of genetic models, we can look at the quantitative contribution of a single locus with genotype
2844:
2817:
377:
A particularly important component of the genetic variance is the additive variance, Var(A), which is the variance due to the average effects (additive effects) of the
3687:
3104:
2899:
Experiments can be run with a similar setup to the one given in Table 1. Using different relationship groups, we can evaluate different intraclass correlations. Using
141:
activity of individual genes associated with environmental changes. However, there are a large number of genes whose transcription is not affected by the environment.
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3159:
3054:
2874:
2436:
269:
5774:
2951:
2924:
2238:
2211:
1852:
3999:
2289:
3079:
5885:
3429:
3314:
3260:
3206:
3185:
3138:
3034:
2894:
2457:
2310:
2184:
2164:
1879:
3514:
When a large, complex pedigree or another aforementioned type of data is available, heritability and other quantitative genetic parameters can be estimated by
5051:
Heckerman D, Gurdasani D, Kadie C, Pomilla C, Carstensen T, Martin H, Ekoru K, Nsubuga RN, Ssenyomo G, Kamali A, Kaleebu P, Widmer C, Sandhu MS (July 2016).
1529:
in which the similarity of identical and fraternal twins is used to estimate heritability. These studies can be limited by the fact that identical twins are
129:
The extent of dependence of phenotype on environment can also be a function of the genes involved. Matters of heritability are complicated because genes may
1961:). The variance will include terms for genetic variance (since they did not all get the same genotype) and environmental variance. This is thought of as an
3944:
3866:
emphasize that heritability is itself a function of environmental variation. However, some researchers argue that it is possible to disentangle the two.
1502:
of breeding studies, using the intraclass correlation of relatives. Various methods of estimating components of variance (and, hence, heritability) from
1113:
844:
1957:
Consider the experiment above. We have two groups of progeny we can compare. The first is comparing the various progeny for an individual sire (called
4422:
3507:
to estimate the influence on a trait by genetic factors, which is reflected by the rate and influence of putatively associated genetic loci (usually
4829:
Cattell RB (November 1960). "The multiple abstract variance analysis equations and solutions: for nature-nurture research on continuous variables".
4940:"Modeling genetic and environmental factors to increase heritability and ease the identification of candidate genes for birth weight: a twin study"
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6852:
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1703:
Figure 3. Twin concordances for seven psychological traits (sample size shown inside bars), with DZ being fraternal and MZ being identical twins.
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1976:
as in the within sire groups, we have an addition term due to the differences among different means of half sibs. The intraclass correlation is
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6178:
6036:
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based on the variance captured by common genetic variants. There are multiple methods that make different adjustments for allele frequency and
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5681:
5637:
5327:
5285:
5206:
5179:
4580:
3975:
1651:'s (1889) data showing the relationship between offspring height (928 individuals) as a function of mean parent height (205 sets of parents).
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6732:
392:
277:
5333:
5291:
6666:
5767:
3700:
Observing the response to selection in an artificial selection experiment will allow calculation of realized heritability as in Fig. 4.
2316:
6239:
358:
is the broad-sense heritability. This reflects all the genetic contributions to a population's phenotypic variance including additive,
6511:
4922:
4638:
3874:
1973:
1514:
6211:
5878:
6254:
6249:
2703:
2682:
5519:
161:. Environment and genetics may also interact, and heritability analyses can test for and examine these interactions (GxE models).
6468:
5973:
3508:
3504:
1510:
1495:
137:, which makes heritability difficult to measure in some cases. Recent insights in molecular biology have identified changes in
4253:
34:
in height between people. This is not the same as asking to what extent do genetic factors influence height in any one person.
6929:
5978:
5760:
5743:
6645:
1669:
1264:
995:
6924:
6496:
4700:
3850:
3515:
5908:
3706:
3548:
2127:{\displaystyle \mathrm {corr} (z,z')=\mathrm {corr} (\mu +{\frac {1}{2}}g+e,\mu +{\frac {1}{2}}g+e')={\frac {1}{4}}V_{g}}
1566:
6934:
5871:
3882:
3545:
1541:
6463:
5953:
4782:"Nurture net of nature: Re-evaluating the role of shared environments in academic achievement and verbal intelligence"
1890:
1624:
1426:
1425:, siblings, parents and offspring, rather than from more distantly related (and therefore less similar) subjects. The
1437:
is generally not possible when gathering human data, relying on naturally occurring relationships and environments.
521:
The simplest genetic model involves a single locus with two alleles (b and B) affecting one quantitative phenotype.
6655:
6521:
6353:
1661:
1472:
26:
6074:
4330:
Maccoby EE (February 2000). "Parenting and its effects on children: on reading and misreading behavior genetics".
1540:, or because of evocative effects (where a genome evokes environments by its effect on them), G and E may covary:
6903:
6348:
5948:
5821:
5360:
5319:
1418:
78:
3858:
contributions of genes and environment, as such traits result from multiple causes interacting. In particular,
517:
Figure 1. Relationship of phenotypic values to additive and dominance effects using a completely dominant locus.
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6126:
5811:
3932:
2463:
138:
1754:
The second set of methods of estimation of heritability involves ANOVA and estimation of variance components.
6883:
6589:
6526:
6501:
6384:
6304:
5938:
3542:
1487:
1440:
In classical quantitative genetics, there were two schools of thought regarding estimation of heritability.
130:
5154:
2964:
6574:
6473:
6458:
6427:
6328:
5923:
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3613:
of plants and animals, the expected response to selection of a trait with known narrow-sense heritability
3586:
3569:
2553:
359:
4448:"The paradox of intelligence: Heritability and malleability coexist in hidden gene-environment interplay"
4383:
3601:
Figure 4. Strength of selection (S) and response to selection (R) in an artificial selection experiment,
2518:
1708:
heritability". Fraternal or dizygotic (DZ) twins on average share half their genes (assuming there is no
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6685:
6579:
6506:
6405:
6343:
6338:
6269:
6204:
6165:
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5958:
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1775:
1491:
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1460:
1452:
134:
90:
4938:
Gielen M, Lindsey PJ, Derom C, Smeets HJ, Souren NY, Paulussen AD, Derom R, Nijhuis JG (January 2008).
1717:
5741:
Quantitative
Genetics Resources website, including the two volume book by Lynch and Walsh. Free access
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1434:
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1390:
502:
145:
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Johnson W, Penke L, Spinath FM (2011). "Understanding
Heritability: What it is and What it is Not".
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The
Trouble with Twin Studies: A Reassessment of Twin Research in the Social and Behavioral Sciences
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1537:
1444:
82:
5053:"Linear mixed model for heritability estimation that explicitly addresses environmental variation"
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For a model with additive and dominance terms, but not others, the equation for a single locus is
513:
6379:
6145:
6141:
6001:
5716:
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5019:
4889:
4630:
4520:
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3911:
3610:
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1709:
94:
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6136:
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5996:
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5708:
5677:
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5605:
5503:
5453:
5404:
5323:
5313:
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5271:
5267:
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5202:
5175:
5143:
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5011:
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4918:
4881:
4846:
4811:
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4673:
4622:
4576:
4512:
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4406:
4347:
4298:
4237:
4166:
4089:
4044:
3981:
3971:
3805:
3656:
3573:
3561:
3086:
2686:
1464:
1377:
490:
182:
173:. In practice, all human behavioral traits vary and almost all traits show some heritability.
114:
98:
66:
3616:
3144:
3039:
6847:
6650:
6619:
6614:
6569:
6197:
6095:
6006:
5700:
5597:
5562:
5495:
5443:
5435:
5396:
5238:
5169:
5133:
5123:
5082:
5072:
5003:
4959:
4951:
4873:
4838:
4801:
4793:
4752:
4742:
4614:
4553:
4504:
4467:
4459:
4398:
4339:
4290:
4227:
4219:
4156:
4148:
4081:
4034:
4026:
3565:
2849:
2406:
382:
244:
205:
Likewise the phenotypic variance in the trait – Var (P) – is the sum of effects as follows:
58:
46:
4599:
3793:
2929:
2902:
2216:
2189:
1830:
6821:
6801:
6660:
6564:
6374:
5913:
5747:
4379:
4134:"High-definition likelihood inference of genetic correlations across human complex traits"
3842:
3834:
2954:
1968:
The second group of progeny are comparisons of means of half sibs with each other (called
483:
367:
30:
Studies of heritability ask questions such as to what extent do genetic factors influence
2268:
1552:
The first school of estimation uses regression and correlation to estimate heritability.
5491:
5428:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
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5111:
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4757:
4723:"The mystery of missing heritability: Genetic interactions create phantom heritability"
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3019:
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2149:
1864:
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The use of ANOVA to calculate heritability often fails to account for the presence of
113:
Heritability measures the fraction of phenotype variability that can be attributed to
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6410:
6244:
6221:
6100:
6090:
6046:
5196:
5023:
4908:
4864:
DeFries JC, Fulker DW (September 1985). "Multiple regression analysis of twin data".
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4524:
4508:
4178:
1483:
1476:
1448:
835:
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4359:
4249:
370:, where individuals are directly affected by their parents' phenotype, such as with
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6274:
6116:
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4101:
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5752:
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4418:
4207:
1251:{\displaystyle \mathrm {Var} (D)=f(bb)d_{bb}^{2}+f(Bb)d_{Bb}^{2}+f(BB)d_{BB}^{2},}
982:{\displaystyle \mathrm {Var} (A)=f(bb)a_{bb}^{2}+f(Bb)a_{Bb}^{2}+f(BB)a_{BB}^{2},}
5566:
5128:
4343:
6816:
6811:
6389:
6369:
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5110:
Hill WG, Goddard ME, Visscher PM (February 2008). MacKay TF, Goddard ME (eds.).
3854:
3826:
1713:
1699:
1668:, the slope is always less than one). This regression effect also underlies the
1643:
6189:
5315:
Doctoring the Mind: Is Our
Current Treatment of Mental Illness Really Any Good?
5057:
Proceedings of the
National Academy of Sciences of the United States of America
4989:"Insensitivity of the analysis of variance to heredity-environment interaction"
4727:
Proceedings of the
National Academy of Sciences of the United States of America
4692:
4294:
4030:
3555:
1742:, reflects the degree to which identical twins raised together are dissimilar,
1520:
Studies of human heritability often utilize adoption study designs, often with
6857:
6741:
6633:
6333:
6286:
5735:
5112:"Data and theory point to mainly additive genetic variance for complex traits"
5007:
4955:
4152:
3830:
1694:
1526:
1471:). It is based on the analysis of correlations and, by extension, regression.
1410:
166:
154:
150:
123:
102:
62:
5712:
5015:
4677:
4626:
4302:
3985:
2671:{\displaystyle H^{2}={\frac {V_{g}}{V_{g}+V_{e}}}={\frac {4(S-W)}{S+(r-1)W}}}
6584:
6552:
6319:
6225:
6121:
5499:
5077:
4747:
4618:
4558:
4539:
363:
158:
42:
16:
Estimation of effect of genetic variation on phenotypic variation of a trait
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5457:
5439:
5408:
5252:
5147:
5096:
4973:
4850:
4815:
4766:
4481:
4410:
4351:
4170:
4093:
4048:
5507:
4885:
4516:
4241:
3869:
The controversy over heritability estimates is largely via their basis in
1107:
There is a similar relationship for the variance of dominance deviations:
6837:
6775:
6547:
6516:
5852:
5842:
5783:
5243:
5226:
4232:
3906:
3523:
1456:
1414:
50:
20:
6279:
4914:
4877:
4463:
4223:
4161:
3551:
3109:
Some common relationships and their coefficients are given in Table 2.
5863:
5736:
Stanford
Encyclopedia of Philosophy entry on Heredity and Heritability
5424:"Missing compared to what? Revisiting heritability, genes and culture"
6259:
5601:
5400:
5277:
4842:
3538:
378:
170:
5704:
4402:
4085:
2953:
as the dominance deviation variance, intraclass correlations become
460:{\displaystyle h^{2}={\frac {\mathrm {Var} (A)}{\mathrm {Var} (P)}}}
345:{\displaystyle H^{2}={\frac {\mathrm {Var} (G)}{\mathrm {Var} (P)}}}
4067:"Concepts, estimation and interpretation of SNP-based heritability"
3534:
The pedigrees can be viewed using programs such as Pedigree Viewer
77:
Other causes of measured variation in a trait are characterized as
5227:"Commentary: heritability estimates--long past their sell-by date"
3596:
2390:{\displaystyle {\frac {3}{4}}V_{g}+V_{e}+r({{\frac {1}{4}}V_{g}})}
1642:
1509:
Today, heritability can be estimated from general pedigrees using
1422:
25:
169:, or because they are omni-present, like if everyone is drinking
6557:
4065:
Yang J, Zeng J, Goddard ME, Wray NR, Visscher PM (August 2017).
1530:
1521:
1429:
for heritability estimates is improved with large sample sizes.
371:
165:
in the population, such as no one having access to a particular
118:
6714:
6193:
5867:
5756:
5540:
Tredoux, Gavan. "The Nature and Nurture of Rectangles." (2019).
4495:
Block N (August 1995). "How heritability misleads about race".
4384:"Heritability in the genomics era--concepts and misconceptions"
1533:, potentially resulting in an underestimation of heritability.
241:) can be controlled and held at 0. In this case, heritability,
5387:
Moore DS, Shenk D (January 2017). "The heritability fallacy".
3776:
2186:
progeny per sire, we can calculate the following ANOVA, using
497:
leads to an estimate of the narrow-sense heritability (called
486:
in which heritability can be estimated and selection modeled.
6710:
4575:. Sunderland, MA: Sinauer and Associates. pp. xv + 652.
3560:
Pedigree models are helpful for untangling confounds such as
2780:{\displaystyle y_{ij}=\mu +\alpha _{i}+\alpha _{j}+d_{ij}+e,}
2689:
for testing for interaction effects than for direct effects.
1672:
for analyzing twins selected for one member being affected.
2138:
since environmental effects are independent of each other.
3113:
Table 2: Coefficients for calculating variance components
3966:
Gazzaniga MS, Heatherton TF, Halpern DF (February 2015).
3703:
Heritability in the above equation is equal to the ratio
1560:
In the comparison of relatives, we find that in general,
4721:
Zuk O, Hechter E, Sunyaev SR, Lander ES (January 2012).
4013:
Yang J, Lee SH, Goddard ME, Visscher PM (January 2011).
3765:
only if the genotype and the environmental noise follow
3801:
5168:
Plomin R, DeFries JC, McClearn GE, McGuffin P (2017).
1367:{\displaystyle f(bb)d_{bb}+f(Bb)d_{Bb}+f(BB)d_{BB}=0.}
1098:{\displaystyle f(bb)a_{bb}+f(Bb)a_{Bb}+f(BB)a_{BB}=0.}
4015:"GCTA: a tool for genome-wide complex trait analysis"
3709:
3659:
3619:
3471:
3443:
3417:
3389:
3356:
3328:
3302:
3274:
3248:
3220:
3194:
3173:
3147:
3126:
3089:
3064:
3042:
3022:
2967:
2932:
2905:
2882:
2852:
2825:
2798:
2706:
2562:
2521:
2466:
2445:
2409:
2319:
2298:
2271:
2219:
2192:
2172:
2152:
1985:
1893:
1867:
1833:
1778:
1569:
1267:
1116:
998:
847:
582:
395:
280:
247:
4672:(1st ed.). Ames, Iowa: Iowa State Univ. Press.
4600:"Three Laws of Behavior Genetics and What They Mean"
2876:
is the dominance deviation for the ij genotype, and
6876:
6830:
6784:
6748:
6678:
6607:
6535:
6489:
6482:
6446:
6398:
6362:
6295:
6232:
6109:
6083:
6045:
6020:
5987:
5901:
5830:
5794:
3825:Heritability estimates' prominent critics, such as
3758:{\displaystyle \mathrm {Var} (A)/\mathrm {Var} (P)}
1609:{\displaystyle h^{2}={\frac {b}{r}}={\frac {t}{r}}}
834:The additive genetic variance at this locus is the
5389:Wiley Interdisciplinary Reviews: Cognitive Science
4663:
4661:
4659:
4373:
4371:
4369:
3757:
3681:
3638:
3484:
3456:
3423:
3402:
3369:
3341:
3308:
3287:
3254:
3233:
3200:
3179:
3153:
3132:
3098:
3073:
3048:
3028:
3005:
2945:
2918:
2888:
2868:
2838:
2811:
2779:
2670:
2544:
2502:
2451:
2430:
2389:
2304:
2283:
2232:
2205:
2178:
2158:
2126:
1941:
1873:
1846:
1816:
1608:
1366:
1250:
1097:
981:
824:
459:
344:
263:
5548:
5546:
3576:, shared environment, and maternal gene effects.
185:as the sum of genetic and environmental effects:
4540:"Principles of Population Genetics, 4th edition"
1942:{\displaystyle z_{i}=\mu +{\frac {1}{2}}g_{i}+e}
74:explained by the environment or random chance?"
5474:Marcus W. Feldman; Richard C. Lewontin (1975).
1498:, as well as other schools. It is based on the
1380:of phenotype on genotype is shown in Figure 1.
381:. Since each parent passes a single allele per
528:alleles can be 0, 1, or 2. For any genotype, (
474:is used to denote broad sense, and lower case
6726:
6205:
5879:
5768:
3837:, focus largely on heritability estimates in
1660:," since the offspring values always tend to
8:
5674:Genetics and analysis of quantitative traits
1656:the slope. (This is the source of the term "
1548:Regression/correlation methods of estimation
5588:Turkheimer E (2015). "Genetic Prediction".
4607:Current Directions in Psychological Science
2556:between half sibs. We can easily calculate
6733:
6719:
6711:
6486:
6212:
6198:
6190:
5886:
5872:
5864:
5775:
5761:
5753:
3998:: CS1 maint: location missing publisher (
1750:Analysis of variance methods of estimation
5447:
5422:Feldman MW, Ramachandran S (April 2018).
5242:
5137:
5127:
5086:
5076:
4963:
4805:
4756:
4746:
4557:
4471:
4231:
4160:
4038:
3735:
3730:
3710:
3708:
3670:
3658:
3627:
3618:
3472:
3470:
3444:
3442:
3416:
3390:
3388:
3357:
3355:
3329:
3327:
3301:
3275:
3273:
3247:
3221:
3219:
3193:
3172:
3146:
3125:
3088:
3063:
3041:
3021:
2994:
2978:
2966:
2937:
2931:
2910:
2904:
2881:
2857:
2851:
2830:
2824:
2803:
2797:
2759:
2746:
2733:
2711:
2705:
2618:
2606:
2593:
2582:
2576:
2567:
2561:
2536:
2522:
2520:
2503:{\displaystyle {\frac {3}{4}}V_{g}+V_{e}}
2494:
2481:
2467:
2465:
2444:
2408:
2377:
2363:
2362:
2347:
2334:
2320:
2318:
2297:
2270:
2224:
2218:
2197:
2191:
2171:
2151:
2118:
2104:
2074:
2046:
2023:
1986:
1984:
1927:
1913:
1898:
1892:
1866:
1838:
1832:
1802:
1783:
1777:
1596:
1583:
1574:
1568:
1349:
1318:
1287:
1266:
1239:
1231:
1203:
1195:
1167:
1159:
1117:
1115:
1080:
1049:
1018:
997:
970:
962:
934:
926:
898:
890:
848:
846:
803:
793:
771:
759:
744:
731:
709:
697:
689:
670:
660:
652:
637:
624:
616:
591:
583:
581:
434:
412:
409:
400:
394:
319:
297:
294:
285:
279:
252:
246:
5174:(2nd ed.). New York: W.H. Freeman.
4907:Falconer DS, Mackay TF (December 1995).
3111:
2846:is the additive effect of the j allele,
2819:is the additive effect of the i allele,
2242:
1698:
838:of the squares of the additive effects:
512:
6899:Suppressed research in the Soviet Union
6853:Inheritance of acquired characteristics
4132:Ning Z, Pawitan Y, Shen X (June 2020).
3923:
2693:Model with additive and dominance terms
1482:The second was originally developed by
366:(multi-genic interactions), as well as
6702:Index of evolutionary biology articles
6179:Index of evolutionary biology articles
5553:Turkheimer E (2011). "Still missing".
5368:from the original on 28 September 2011
3991:
5231:International Journal of Epidemiology
5198:Introduction to quantitative genetics
4910:Introduction to Quantitative Genetics
4780:Daw J, Guo G, Harris KM (July 2015).
4670:An introduction to genetic statistics
3537:, and analyzed with programs such as
3006:{\displaystyle =rV_{a}+\theta V_{d},}
2926:as the additive genetic variance and
1631:is the coefficient of regression and
1525:second and more common design is the
1479:as a way of estimating heritability.
7:
6889:Collectivization in the Soviet Union
5676:. Sunderland, Mass.: Sinauer Assoc.
4644:from the original on 19 October 2013
4127:
4125:
4060:
4058:
3547:, MCMCglmm within the R environment
1531:not completely genetically identical
501:). This is the principle underlying
5632:. New York: Routledge. p. 81.
4697:Stanford Encyclopedia of Philosophy
4691:Stephen Downes and Lucas Matthews.
4446:Sauce B, Matzel LD (January 2018).
4208:"Coming to terms with heritability"
2545:{\displaystyle {\frac {1}{4}}V_{g}}
2244:Table 1: ANOVA for Sire experiment
1635:is the coefficient of correlation.
489:Additive variance is important for
6512:Evolutionary developmental biology
4276:"The intelligence of heritability"
4019:American Journal of Human Genetics
3947:from the original on 2 August 2015
3742:
3739:
3736:
3717:
3714:
3711:
2033:
2030:
2027:
2024:
1996:
1993:
1990:
1987:
1817:{\displaystyle y_{i}=\mu +g_{i}+e}
1726:The effect of shared environment,
1413:analyses required to estimate the
1124:
1121:
1118:
855:
852:
849:
441:
438:
435:
419:
416:
413:
326:
323:
320:
304:
301:
298:
14:
5795:Concepts in Quantitative Genetics
4573:Principles of Population Genetics
2957:of these parameters. In general,
2685:, because ANOVA has a much lower
1738:. Unique environmental variance,
6469:Evolution of sexual reproduction
5522:from the original on 20 May 2021
5201:(4th ed.). Essex: Longman.
5153:
4987:Wahlsten, Douglas (March 1990).
4798:10.1016/j.ssresearch.2015.02.011
3892:is a concept widely applicable.
3781:
1517:estimated from genetic markers.
181:Any particular phenotype can be
5693:European Journal of Personality
5649:from the original on 2016-04-04
5336:from the original on 2020-10-05
5294:from the original on 2017-07-19
5195:Falconer DS, Mackay TF (1998).
5033:from the original on 2020-10-05
4703:from the original on 2020-02-25
4428:from the original on 2016-03-24
4312:from the original on 2018-10-24
4256:from the original on 2020-12-02
4188:from the original on 2021-04-15
4111:from the original on 2020-10-05
3933:"Estimating Trait Heritability"
3509:single-nucleotide polymorphisms
3505:genome-wide association studies
3485:{\displaystyle {\frac {1}{16}}}
2240:as the environmental variance:
1496:North Carolina State University
386:heritability and is defined as
6240:Genotype–phenotype distinction
5979:Constructive neutral evolution
3752:
3746:
3727:
3721:
3633:
3620:
3457:{\displaystyle {\frac {1}{4}}}
3403:{\displaystyle {\frac {1}{8}}}
3370:{\displaystyle {\frac {1}{4}}}
3342:{\displaystyle {\frac {1}{2}}}
3288:{\displaystyle {\frac {1}{4}}}
3234:{\displaystyle {\frac {1}{2}}}
2683:gene–-environment interactions
2659:
2647:
2636:
2624:
2425:
2413:
2384:
2359:
2098:
2037:
2017:
2000:
1716:between MZ and DZ twins, i.e.
1342:
1333:
1311:
1302:
1280:
1271:
1224:
1215:
1188:
1179:
1152:
1143:
1134:
1128:
1073:
1064:
1042:
1033:
1011:
1002:
955:
946:
919:
910:
883:
874:
865:
859:
812:
809:
786:
764:
753:
750:
724:
702:
676:
653:
643:
617:
451:
445:
429:
423:
336:
330:
314:
308:
144:Estimates of heritability use
1:
6497:Regulation of gene expression
5555:Research in Human Development
5171:Behavioral Genetics: A Primer
4996:Behavioral and Brain Sciences
4598:Turkheimer E (October 2000).
3516:restricted maximum likelihood
1881:is the environmental effect.
1556:Comparison of close relatives
1455:, and further popularized by
368:maternal and paternal effects
53:that estimates the degree of
6667:Endless Forms Most Beautiful
6447:Evolution of genetic systems
6255:Gene–environment correlation
6250:Gene–environment interaction
5929:Fisher's fundamental theorem
5567:10.1080/15427609.2011.625321
5356:"Doctoring the Mind: Review"
5129:10.1371/journal.pgen.1000008
4509:10.1016/0010-0277(95)00678-r
4344:10.1146/annurev.psych.51.1.1
4206:Stoltenberg SF (June 1997).
3881:studies' conclusions is the
3877:studies to corroborate such
3804:or discuss the issue on the
2213:as the genetic variance and
1542:gene environment correlation
1506:are used in these analyses.
233:In a planned experiment Cov(
6646:Christiane NĂĽsslein-Volhard
5954:Coefficient of relationship
4571:Hartl DL, Clark AG (2007).
4332:Annual Review of Psychology
3931:Wray N, Visscher P (2008).
3646:can be estimated using the
2839:{\displaystyle \alpha _{j}}
2812:{\displaystyle \alpha _{i}}
1639:Parent-offspring regression
1488:The University of Edinburgh
6953:
6522:Hedgehog signaling pathway
6399:Developmental architecture
5590:The Hastings Center Report
5476:"The Heritability Hang–Up"
4295:10.1037/0708-5591.35.3.244
4031:10.1016/j.ajhg.2010.11.011
3970:(5th ed.). New York.
3796:towards certain viewpoints
1854:is the effect of genotype
1692:
1664:value for the population,
1625:coefficient of relatedness
18:
6904:Politicization of science
6699:
6349:Transgressive segregation
6174:
5949:Coefficient of inbreeding
5822:Effective population size
5672:Lynch M, Walsh B (1998).
5354:McGrath M (5 July 2009).
5320:New York University Press
5008:10.1017/S0140525X00077797
4956:10.1007/s10519-007-9170-3
4153:10.1038/s41588-020-0653-y
3873:. The scarce success of
1623:can be thought of as the
1453:The University of Chicago
761:Dominance Deviation
484:liability threshold model
6127:Evolutionary game theory
5909:Hardy–Weinberg principle
5812:Quantitative trait locus
4391:Nature Reviews. Genetics
4382:, Wray NR (April 2008).
3682:{\displaystyle R=h^{2}S}
3099:{\displaystyle \theta =}
19:Not to be confused with
6884:Bourgeois pseudoscience
6527:Notch signaling pathway
6502:Gene regulatory network
6385:Dual inheritance theory
5939:Shifting balance theory
5500:10.1126/science.1198102
5078:10.1073/pnas.1510497113
4786:Social Science Research
4748:10.1073/pnas.1119675109
4699:. Stanford University.
4619:10.1111/1467-8721.00084
3639:{\displaystyle (h^{2})}
3154:{\displaystyle \theta }
3049:{\displaystyle \theta }
1953:Intraclass correlations
1401:Estimating heritability
374:production in mammals.
6575:cis-regulatory element
6483:Control of development
6363:Non-genetic influences
6329:evolutionary landscape
5924:Linkage disequilibrium
5440:10.1098/rstb.2017.0064
4452:Psychological Bulletin
3767:Gaussian distributions
3759:
3683:
3640:
3606:
3587:linkage disequilibrium
3486:
3458:
3425:
3404:
3371:
3343:
3310:
3289:
3256:
3235:
3202:
3181:
3155:
3134:
3100:
3075:
3050:
3030:
3007:
2961:Intraclass correlation
2947:
2920:
2890:
2870:
2869:{\displaystyle d_{ij}}
2840:
2813:
2781:
2672:
2554:intraclass correlation
2546:
2504:
2453:
2432:
2431:{\displaystyle n(r-1)}
2391:
2306:
2285:
2234:
2207:
2180:
2160:
2146:In an experiment with
2128:
1972:). In addition to the
1943:
1875:
1848:
1818:
1704:
1652:
1610:
1368:
1252:
1099:
983:
826:
518:
461:
346:
265:
264:{\displaystyle H^{2},}
45:used in the fields of
35:
6930:Quantitative genetics
6863:Mendelian inheritance
6686:Nature versus nurture
6590:Cell surface receptor
6507:Evo-devo gene toolkit
6406:Developmental biology
6344:Polygenic inheritance
6270:Quantitative genetics
6166:Quantitative genetics
6075:Balding–Nichols model
6060:Population bottleneck
6055:Small population size
5959:Selection coefficient
5788:Quantitative genetics
4668:Kempthorne O (1957).
4559:10.1093/jhered/esm035
3968:Psychological science
3847:genetic determination
3760:
3684:
3641:
3600:
3593:Response to selection
3572:, and confounding of
3487:
3459:
3435:Double First Cousins
3426:
3405:
3372:
3344:
3311:
3290:
3257:
3236:
3203:
3182:
3156:
3135:
3101:
3076:
3051:
3031:
3008:
2948:
2946:{\displaystyle V_{d}}
2921:
2919:{\displaystyle V_{a}}
2891:
2871:
2841:
2814:
2782:
2673:
2547:
2505:
2454:
2433:
2392:
2307:
2286:
2258:Expected Mean Square
2235:
2233:{\displaystyle V_{e}}
2208:
2206:{\displaystyle V_{g}}
2181:
2161:
2129:
1944:
1876:
1849:
1847:{\displaystyle g_{i}}
1819:
1702:
1670:DeFries–Fulker method
1646:
1611:
1538:observational studies
1492:Iowa State University
1469:Iowa State University
1461:University of Chicago
1369:
1253:
1100:
984:
827:
699:Additive Effect
516:
499:realized heritability
495:response to selection
462:
347:
266:
135:phenotypic plasticity
91:quantitative genetics
89:important concept in
79:environmental factors
29:
6925:Genetic epidemiology
6595:Transcription factor
6310:Genetic assimilation
6297:Genetic architecture
6037:Background selection
6024:on genomic variation
6022:Effects of selection
5974:Population structure
5848:Evolutionary biology
5322:. pp. 123–127.
4831:Psychological Review
3884:missing heritability
3707:
3657:
3617:
3580:Genomic heritability
3570:prenatal environment
3469:
3441:
3415:
3387:
3354:
3326:
3300:
3272:
3246:
3218:
3192:
3171:
3145:
3124:
3087:
3062:
3040:
3020:
2965:
2930:
2903:
2896:is the environment.
2880:
2850:
2823:
2796:
2704:
2560:
2519:
2464:
2443:
2407:
2317:
2296:
2269:
2263:Between sire groups
2217:
2190:
2170:
2150:
1983:
1891:
1865:
1831:
1776:
1567:
1500:analysis of variance
1435:experimental control
1395:academic achievement
1391:missing heritability
1265:
1114:
996:
845:
580:
503:artificial selection
393:
278:
245:
146:statistical analyses
6935:Population genetics
6797:Georgii Karpechenko
6691:Morphogenetic field
6608:Influential figures
6156:Population genomics
6032:Genetic hitchhiking
5919:Identity by descent
5895:Population genetics
5838:Population genetics
5492:1975Sci...190.1163F
5486:(4220): 1163–1168.
5312:Bentall RP (2009).
5069:2016PNAS..113.7377H
4739:2012PNAS..109.1193Z
4545:Journal of Heredity
4283:Canadian Psychology
4274:Wahlsten D (1994).
3902:Behavioral genetics
3839:behavioral sciences
3802:improve the article
3554:family of programs
3498:Linear mixed models
3114:
2401:Within sire groups
2284:{\displaystyle n-1}
2245:
1662:regress to the mean
1515:genomic relatedness
1511:linear mixed models
1244:
1208:
1172:
975:
939:
903:
83:observational error
6380:Genomic imprinting
6142:Landscape genetics
5746:2006-02-06 at the
5596:(5 Suppl): S32–8.
5434:(1743): 20170064.
5244:10.1093/ije/dyl064
4878:10.1007/BF01066239
4464:10.1037/bul0000131
4224:10.1007/BF02259512
3912:Heritability of IQ
3879:population-genetic
3755:
3679:
3648:breeder's equation
3636:
3611:selective breeding
3607:
3482:
3454:
3421:
3400:
3367:
3339:
3306:
3285:
3252:
3231:
3198:
3177:
3151:
3130:
3112:
3096:
3074:{\displaystyle r=}
3071:
3046:
3026:
3003:
2943:
2916:
2886:
2866:
2836:
2809:
2777:
2668:
2542:
2500:
2449:
2428:
2387:
2302:
2281:
2243:
2230:
2203:
2176:
2156:
2124:
1939:
1871:
1844:
1814:
1718:Falconer's formula
1710:assortative mating
1705:
1676:Sibling comparison
1653:
1606:
1364:
1248:
1227:
1191:
1155:
1095:
979:
958:
922:
886:
822:
820:
519:
478:for narrow sense.
457:
342:
261:
95:selective breeding
93:, particularly in
36:
6912:
6911:
6894:Pavlovian session
6766:Nikita Khrushchev
6708:
6707:
6641:Eric F. Wieschaus
6603:
6602:
6421:Pattern formation
6325:Fitness landscape
6187:
6186:
6137:Genetic genealogy
6132:Fitness landscape
5861:
5860:
5683:978-0-87893-481-2
5639:978-1-317-60590-4
5623:Joseph J (2014).
5329:978-0-8147-8723-6
5287:978-1-898059-47-9
5273:The Gene Illusion
5266:Joseph J (2004).
5208:978-0-582-24302-6
5181:978-0-7167-2056-0
4944:Behavior Genetics
4866:Behavior Genetics
4582:978-0-87893-308-2
3977:978-0-393-26313-8
3875:molecular-genetic
3823:
3822:
3574:genetic dominance
3562:reverse causality
3495:
3494:
3480:
3452:
3424:{\displaystyle 0}
3398:
3365:
3337:
3309:{\displaystyle 0}
3283:
3255:{\displaystyle 0}
3229:
3212:Parent-Offspring
3201:{\displaystyle 1}
3180:{\displaystyle 1}
3133:{\displaystyle r}
3029:{\displaystyle r}
2889:{\displaystyle e}
2687:statistical power
2666:
2613:
2530:
2513:
2512:
2475:
2452:{\displaystyle W}
2371:
2328:
2305:{\displaystyle S}
2179:{\displaystyle r}
2159:{\displaystyle n}
2112:
2082:
2054:
1959:within sire group
1921:
1874:{\displaystyle e}
1723:=2(r(MZ)-r(DZ)).
1604:
1591:
1475:was developed by
1447:was developed by
1445:school of thought
1378:linear regression
762:
700:
692:
455:
340:
197:) + Environment (
115:genetic variation
99:behavior genetics
67:genetic variation
6942:
6735:
6728:
6721:
6712:
6651:William McGinnis
6620:Richard Lewontin
6615:C. H. Waddington
6487:
6464:Neutral networks
6214:
6207:
6200:
6191:
6096:J. B. S. Haldane
5888:
5881:
5874:
5865:
5777:
5770:
5763:
5754:
5724:
5687:
5658:
5657:
5655:
5654:
5648:
5631:
5620:
5614:
5613:
5602:10.1002/hast.496
5585:
5579:
5578:
5561:(3–4): 227–241.
5550:
5541:
5538:
5532:
5531:
5529:
5527:
5471:
5465:
5464:
5451:
5419:
5413:
5412:
5401:10.1002/wcs.1400
5384:
5378:
5377:
5375:
5373:
5351:
5345:
5344:
5342:
5341:
5309:
5303:
5302:
5300:
5299:
5263:
5257:
5256:
5246:
5219:
5213:
5212:
5192:
5186:
5185:
5165:
5159:
5158:
5157:
5151:
5141:
5131:
5107:
5101:
5100:
5090:
5080:
5048:
5042:
5041:
5039:
5038:
5032:
4993:
4984:
4978:
4977:
4967:
4935:
4929:
4928:
4913:(4th ed.).
4904:
4898:
4897:
4861:
4855:
4854:
4843:10.1037/h0043487
4826:
4820:
4819:
4809:
4777:
4771:
4770:
4760:
4750:
4718:
4712:
4711:
4709:
4708:
4688:
4682:
4681:
4665:
4654:
4653:
4651:
4649:
4643:
4604:
4595:
4589:
4586:
4563:
4561:
4538:Wills C (2007).
4535:
4529:
4528:
4492:
4486:
4485:
4475:
4443:
4437:
4436:
4434:
4433:
4427:
4388:
4375:
4364:
4363:
4327:
4321:
4320:
4318:
4317:
4311:
4280:
4271:
4265:
4264:
4262:
4261:
4235:
4203:
4197:
4196:
4194:
4193:
4187:
4164:
4138:
4129:
4120:
4119:
4117:
4116:
4110:
4080:(9): 1304–1310.
4071:
4062:
4053:
4052:
4042:
4010:
4004:
4003:
3997:
3989:
3963:
3957:
3956:
3954:
3952:
3937:Nature Education
3928:
3818:
3815:
3809:
3785:
3784:
3777:
3764:
3762:
3761:
3756:
3745:
3734:
3720:
3688:
3686:
3685:
3680:
3675:
3674:
3645:
3643:
3642:
3637:
3632:
3631:
3566:maternal effects
3520:Bayesian methods
3491:
3489:
3488:
3483:
3481:
3473:
3463:
3461:
3460:
3455:
3453:
3445:
3430:
3428:
3427:
3422:
3409:
3407:
3406:
3401:
3399:
3391:
3376:
3374:
3373:
3368:
3366:
3358:
3348:
3346:
3345:
3340:
3338:
3330:
3315:
3313:
3312:
3307:
3294:
3292:
3291:
3286:
3284:
3276:
3261:
3259:
3258:
3253:
3240:
3238:
3237:
3232:
3230:
3222:
3207:
3205:
3204:
3199:
3186:
3184:
3183:
3178:
3165:Identical Twins
3160:
3158:
3157:
3152:
3139:
3137:
3136:
3131:
3115:
3105:
3103:
3102:
3097:
3080:
3078:
3077:
3072:
3055:
3053:
3052:
3047:
3035:
3033:
3032:
3027:
3012:
3010:
3009:
3004:
2999:
2998:
2983:
2982:
2955:linear functions
2952:
2950:
2949:
2944:
2942:
2941:
2925:
2923:
2922:
2917:
2915:
2914:
2895:
2893:
2892:
2887:
2875:
2873:
2872:
2867:
2865:
2864:
2845:
2843:
2842:
2837:
2835:
2834:
2818:
2816:
2815:
2810:
2808:
2807:
2786:
2784:
2783:
2778:
2767:
2766:
2751:
2750:
2738:
2737:
2719:
2718:
2677:
2675:
2674:
2669:
2667:
2665:
2639:
2619:
2614:
2612:
2611:
2610:
2598:
2597:
2587:
2586:
2577:
2572:
2571:
2551:
2549:
2548:
2543:
2541:
2540:
2531:
2523:
2509:
2507:
2506:
2501:
2499:
2498:
2486:
2485:
2476:
2468:
2458:
2456:
2455:
2450:
2437:
2435:
2434:
2429:
2396:
2394:
2393:
2388:
2383:
2382:
2381:
2372:
2364:
2352:
2351:
2339:
2338:
2329:
2321:
2311:
2309:
2308:
2303:
2290:
2288:
2287:
2282:
2246:
2239:
2237:
2236:
2231:
2229:
2228:
2212:
2210:
2209:
2204:
2202:
2201:
2185:
2183:
2182:
2177:
2165:
2163:
2162:
2157:
2133:
2131:
2130:
2125:
2123:
2122:
2113:
2105:
2097:
2083:
2075:
2055:
2047:
2036:
2016:
1999:
1970:among sire group
1948:
1946:
1945:
1940:
1932:
1931:
1922:
1914:
1903:
1902:
1880:
1878:
1877:
1872:
1853:
1851:
1850:
1845:
1843:
1842:
1823:
1821:
1820:
1815:
1807:
1806:
1788:
1787:
1615:
1613:
1612:
1607:
1605:
1597:
1592:
1584:
1579:
1578:
1486:and expanded at
1373:
1371:
1370:
1365:
1357:
1356:
1326:
1325:
1295:
1294:
1257:
1255:
1254:
1249:
1243:
1238:
1207:
1202:
1171:
1166:
1127:
1104:
1102:
1101:
1096:
1088:
1087:
1057:
1056:
1026:
1025:
988:
986:
985:
980:
974:
969:
938:
933:
902:
897:
858:
836:weighted average
831:
829:
828:
823:
821:
808:
807:
798:
797:
779:
778:
763:
760:
749:
748:
736:
735:
717:
716:
701:
698:
693:
690:
682:
675:
674:
665:
664:
642:
641:
629:
628:
599:
598:
466:
464:
463:
458:
456:
454:
444:
432:
422:
410:
405:
404:
351:
349:
348:
343:
341:
339:
329:
317:
307:
295:
290:
289:
270:
268:
267:
262:
257:
256:
59:phenotypic trait
6952:
6951:
6945:
6944:
6943:
6941:
6940:
6939:
6915:
6914:
6913:
6908:
6877:Soviet policies
6872:
6826:
6822:Nikolai Vavilov
6802:Zhores Medvedev
6792:Wacław Gajewski
6780:
6744:
6739:
6709:
6704:
6695:
6674:
6661:Sean B. Carroll
6599:
6531:
6478:
6442:
6394:
6375:Maternal effect
6358:
6291:
6228:
6218:
6188:
6183:
6170:
6105:
6079:
6041:
6025:
6023:
6016:
5983:
5914:Genetic linkage
5897:
5892:
5862:
5857:
5826:
5790:
5781:
5748:Wayback Machine
5732:
5727:
5705:10.1002/per.835
5690:
5684:
5671:
5667:
5665:Further reading
5662:
5661:
5652:
5650:
5646:
5640:
5629:
5622:
5621:
5617:
5587:
5586:
5582:
5552:
5551:
5544:
5539:
5535:
5525:
5523:
5473:
5472:
5468:
5421:
5420:
5416:
5386:
5385:
5381:
5371:
5369:
5353:
5352:
5348:
5339:
5337:
5330:
5311:
5310:
5306:
5297:
5295:
5288:
5280:. p. 141.
5265:
5264:
5260:
5221:
5220:
5216:
5209:
5194:
5193:
5189:
5182:
5167:
5166:
5162:
5152:
5122:(2): e1000008.
5109:
5108:
5104:
5063:(27): 7377–82.
5050:
5049:
5045:
5036:
5034:
5030:
4991:
4986:
4985:
4981:
4937:
4936:
4932:
4925:
4906:
4905:
4901:
4863:
4862:
4858:
4828:
4827:
4823:
4779:
4778:
4774:
4720:
4719:
4715:
4706:
4704:
4690:
4689:
4685:
4667:
4666:
4657:
4647:
4645:
4641:
4602:
4597:
4596:
4592:
4583:
4570:
4548:(Book Review).
4537:
4536:
4532:
4494:
4493:
4489:
4445:
4444:
4440:
4431:
4429:
4425:
4403:10.1038/nrg2322
4386:
4377:
4376:
4367:
4329:
4328:
4324:
4315:
4313:
4309:
4278:
4273:
4272:
4268:
4259:
4257:
4205:
4204:
4200:
4191:
4189:
4185:
4141:Nature Genetics
4136:
4131:
4130:
4123:
4114:
4112:
4108:
4086:10.1038/ng.3941
4074:Nature Genetics
4069:
4064:
4063:
4056:
4012:
4011:
4007:
3990:
3978:
3965:
3964:
3960:
3950:
3948:
3930:
3929:
3925:
3920:
3898:
3843:social sciences
3835:Richard Bentall
3819:
3813:
3810:
3799:
3786:
3782:
3775:
3705:
3704:
3666:
3655:
3654:
3623:
3615:
3614:
3595:
3582:
3500:
3467:
3466:
3439:
3438:
3413:
3412:
3385:
3384:
3352:
3351:
3324:
3323:
3298:
3297:
3270:
3269:
3244:
3243:
3216:
3215:
3190:
3189:
3169:
3168:
3143:
3142:
3122:
3121:
3085:
3084:
3060:
3059:
3038:
3037:
3018:
3017:
2990:
2974:
2963:
2962:
2933:
2928:
2927:
2906:
2901:
2900:
2878:
2877:
2853:
2848:
2847:
2826:
2821:
2820:
2799:
2794:
2793:
2755:
2742:
2729:
2707:
2702:
2701:
2695:
2640:
2620:
2602:
2589:
2588:
2578:
2563:
2558:
2557:
2532:
2517:
2516:
2490:
2477:
2462:
2461:
2441:
2440:
2405:
2404:
2373:
2343:
2330:
2315:
2314:
2294:
2293:
2267:
2266:
2220:
2215:
2214:
2193:
2188:
2187:
2168:
2167:
2148:
2147:
2144:
2114:
2090:
2009:
1981:
1980:
1955:
1923:
1894:
1889:
1888:
1863:
1862:
1859:
1834:
1829:
1828:
1798:
1779:
1774:
1773:
1767:
1760:
1752:
1697:
1691:
1678:
1641:
1570:
1565:
1564:
1558:
1550:
1522:identical twins
1403:
1386:
1345:
1314:
1283:
1263:
1262:
1112:
1111:
1076:
1045:
1014:
994:
993:
843:
842:
819:
818:
799:
789:
767:
740:
727:
705:
691:Population mean
680:
679:
666:
656:
633:
620:
600:
587:
578:
577:
573:
566:
555:
548:
541:
534:
511:
433:
411:
396:
391:
390:
318:
296:
281:
276:
275:
248:
243:
242:
179:
139:transcriptional
111:
101:(for instance,
65:that is due to
24:
17:
12:
11:
5:
6950:
6949:
6946:
6938:
6937:
6932:
6927:
6917:
6916:
6910:
6909:
6907:
6906:
6901:
6896:
6891:
6886:
6880:
6878:
6874:
6873:
6871:
6870:
6865:
6860:
6855:
6850:
6845:
6840:
6834:
6832:
6828:
6827:
6825:
6824:
6819:
6814:
6809:
6807:Georgii Nadson
6804:
6799:
6794:
6788:
6786:
6782:
6781:
6779:
6778:
6773:
6768:
6763:
6758:
6756:Trofim Lysenko
6752:
6750:
6746:
6745:
6740:
6738:
6737:
6730:
6723:
6715:
6706:
6705:
6700:
6697:
6696:
6694:
6693:
6688:
6682:
6680:
6676:
6675:
6673:
6672:
6671:
6670:
6658:
6653:
6648:
6643:
6638:
6637:
6636:
6625:François Jacob
6622:
6617:
6611:
6609:
6605:
6604:
6601:
6600:
6598:
6597:
6592:
6587:
6582:
6577:
6572:
6567:
6562:
6561:
6560:
6550:
6545:
6539:
6537:
6533:
6532:
6530:
6529:
6524:
6519:
6514:
6509:
6504:
6499:
6493:
6491:
6484:
6480:
6479:
6477:
6476:
6471:
6466:
6461:
6456:
6450:
6448:
6444:
6443:
6441:
6440:
6435:
6430:
6425:
6424:
6423:
6418:
6408:
6402:
6400:
6396:
6395:
6393:
6392:
6387:
6382:
6377:
6372:
6366:
6364:
6360:
6359:
6357:
6356:
6354:Sequence space
6351:
6346:
6341:
6336:
6331:
6322:
6317:
6312:
6307:
6301:
6299:
6293:
6292:
6290:
6289:
6284:
6283:
6282:
6272:
6267:
6262:
6257:
6252:
6247:
6242:
6236:
6234:
6230:
6229:
6219:
6217:
6216:
6209:
6202:
6194:
6185:
6184:
6182:
6181:
6175:
6172:
6171:
6169:
6168:
6163:
6161:Phylogeography
6158:
6153:
6151:Microevolution
6148:
6139:
6134:
6129:
6124:
6119:
6113:
6111:
6110:Related topics
6107:
6106:
6104:
6103:
6098:
6093:
6087:
6085:
6081:
6080:
6078:
6077:
6072:
6067:
6065:Founder effect
6062:
6057:
6051:
6049:
6043:
6042:
6040:
6039:
6034:
6028:
6026:
6021:
6018:
6017:
6015:
6014:
6009:
6004:
5999:
5993:
5991:
5985:
5984:
5982:
5981:
5976:
5971:
5966:
5961:
5956:
5951:
5946:
5944:Price equation
5941:
5936:
5934:Neutral theory
5931:
5926:
5921:
5916:
5911:
5905:
5903:
5899:
5898:
5893:
5891:
5890:
5883:
5876:
5868:
5859:
5858:
5856:
5855:
5850:
5845:
5840:
5834:
5832:
5831:Related Topics
5828:
5827:
5825:
5824:
5819:
5817:Candidate gene
5814:
5809:
5804:
5798:
5796:
5792:
5791:
5782:
5780:
5779:
5772:
5765:
5757:
5751:
5750:
5738:
5731:
5730:External links
5728:
5726:
5725:
5699:(4): 287–294.
5688:
5682:
5668:
5666:
5663:
5660:
5659:
5638:
5615:
5580:
5542:
5533:
5466:
5414:
5395:(1–2): e1400.
5379:
5346:
5328:
5304:
5286:
5258:
5214:
5207:
5187:
5180:
5160:
5102:
5043:
5002:(1): 109–120.
4979:
4930:
4924:978-0582243026
4923:
4899:
4856:
4821:
4772:
4713:
4693:"Heritability"
4683:
4655:
4613:(5): 160–164.
4590:
4588:
4587:
4581:
4530:
4487:
4438:
4365:
4322:
4289:(3): 244–260.
4266:
4218:(2–3): 89–96.
4198:
4147:(8): 859–864.
4121:
4054:
4005:
3976:
3958:
3922:
3921:
3919:
3916:
3915:
3914:
3909:
3904:
3897:
3894:
3821:
3820:
3789:
3787:
3780:
3774:
3771:
3754:
3751:
3748:
3744:
3741:
3738:
3733:
3729:
3726:
3723:
3719:
3716:
3713:
3690:
3689:
3678:
3673:
3669:
3665:
3662:
3635:
3630:
3626:
3622:
3594:
3591:
3581:
3578:
3499:
3496:
3493:
3492:
3479:
3476:
3464:
3451:
3448:
3436:
3432:
3431:
3420:
3410:
3397:
3394:
3382:
3381:First Cousins
3378:
3377:
3364:
3361:
3349:
3336:
3333:
3321:
3320:Full Siblings
3317:
3316:
3305:
3295:
3282:
3279:
3267:
3266:Half Siblings
3263:
3262:
3251:
3241:
3228:
3225:
3213:
3209:
3208:
3197:
3187:
3176:
3166:
3162:
3161:
3150:
3140:
3129:
3119:
3095:
3092:
3070:
3067:
3045:
3025:
3014:
3013:
3002:
2997:
2993:
2989:
2986:
2981:
2977:
2973:
2970:
2940:
2936:
2913:
2909:
2885:
2863:
2860:
2856:
2833:
2829:
2806:
2802:
2788:
2787:
2776:
2773:
2770:
2765:
2762:
2758:
2754:
2749:
2745:
2741:
2736:
2732:
2728:
2725:
2722:
2717:
2714:
2710:
2694:
2691:
2664:
2661:
2658:
2655:
2652:
2649:
2646:
2643:
2638:
2635:
2632:
2629:
2626:
2623:
2617:
2609:
2605:
2601:
2596:
2592:
2585:
2581:
2575:
2570:
2566:
2539:
2535:
2529:
2526:
2511:
2510:
2497:
2493:
2489:
2484:
2480:
2474:
2471:
2459:
2448:
2438:
2427:
2424:
2421:
2418:
2415:
2412:
2402:
2398:
2397:
2386:
2380:
2376:
2370:
2367:
2361:
2358:
2355:
2350:
2346:
2342:
2337:
2333:
2327:
2324:
2312:
2301:
2291:
2280:
2277:
2274:
2264:
2260:
2259:
2256:
2253:
2250:
2227:
2223:
2200:
2196:
2175:
2155:
2143:
2140:
2136:
2135:
2121:
2117:
2111:
2108:
2103:
2100:
2096:
2093:
2089:
2086:
2081:
2078:
2073:
2070:
2067:
2064:
2061:
2058:
2053:
2050:
2045:
2042:
2039:
2035:
2032:
2029:
2026:
2022:
2019:
2015:
2012:
2008:
2005:
2002:
1998:
1995:
1992:
1989:
1954:
1951:
1950:
1949:
1938:
1935:
1930:
1926:
1920:
1917:
1912:
1909:
1906:
1901:
1897:
1870:
1857:
1841:
1837:
1825:
1824:
1813:
1810:
1805:
1801:
1797:
1794:
1791:
1786:
1782:
1765:
1759:
1756:
1751:
1748:
1693:Main article:
1690:
1687:
1677:
1674:
1649:Francis Galton
1640:
1637:
1617:
1616:
1603:
1600:
1595:
1590:
1587:
1582:
1577:
1573:
1557:
1554:
1549:
1546:
1427:standard error
1402:
1399:
1385:
1382:
1363:
1360:
1355:
1352:
1348:
1344:
1341:
1338:
1335:
1332:
1329:
1324:
1321:
1317:
1313:
1310:
1307:
1304:
1301:
1298:
1293:
1290:
1286:
1282:
1279:
1276:
1273:
1270:
1259:
1258:
1247:
1242:
1237:
1234:
1230:
1226:
1223:
1220:
1217:
1214:
1211:
1206:
1201:
1198:
1194:
1190:
1187:
1184:
1181:
1178:
1175:
1170:
1165:
1162:
1158:
1154:
1151:
1148:
1145:
1142:
1139:
1136:
1133:
1130:
1126:
1123:
1120:
1094:
1091:
1086:
1083:
1079:
1075:
1072:
1069:
1066:
1063:
1060:
1055:
1052:
1048:
1044:
1041:
1038:
1035:
1032:
1029:
1024:
1021:
1017:
1013:
1010:
1007:
1004:
1001:
990:
989:
978:
973:
968:
965:
961:
957:
954:
951:
948:
945:
942:
937:
932:
929:
925:
921:
918:
915:
912:
909:
906:
901:
896:
893:
889:
885:
882:
879:
876:
873:
870:
867:
864:
861:
857:
854:
851:
817:
814:
811:
806:
802:
796:
792:
788:
785:
782:
777:
774:
770:
766:
758:
755:
752:
747:
743:
739:
734:
730:
726:
723:
720:
715:
712:
708:
704:
696:
688:
685:
683:
681:
678:
673:
669:
663:
659:
655:
651:
648:
645:
640:
636:
632:
627:
623:
619:
615:
612:
609:
606:
603:
601:
597:
594:
590:
586:
585:
571:
564:
553:
546:
539:
532:
524:The number of
510:
507:
470:An upper case
468:
467:
453:
450:
447:
443:
440:
437:
431:
428:
425:
421:
418:
415:
408:
403:
399:
353:
352:
338:
335:
332:
328:
325:
322:
316:
313:
310:
306:
303:
300:
293:
288:
284:
271:is defined as
260:
255:
251:
231:
230:
203:
202:
193:) = Genotype (
178:
175:
110:
107:
15:
13:
10:
9:
6:
4:
3:
2:
6948:
6947:
6936:
6933:
6931:
6928:
6926:
6923:
6922:
6920:
6905:
6902:
6900:
6897:
6895:
6892:
6890:
6887:
6885:
6882:
6881:
6879:
6875:
6869:
6868:Vernalization
6866:
6864:
6861:
6859:
6856:
6854:
6851:
6849:
6848:Hybridization
6846:
6844:
6841:
6839:
6836:
6835:
6833:
6829:
6823:
6820:
6818:
6815:
6813:
6810:
6808:
6805:
6803:
6800:
6798:
6795:
6793:
6790:
6789:
6787:
6783:
6777:
6774:
6772:
6771:Joseph Stalin
6769:
6767:
6764:
6762:
6759:
6757:
6754:
6753:
6751:
6747:
6743:
6736:
6731:
6729:
6724:
6722:
6717:
6716:
6713:
6703:
6698:
6692:
6689:
6687:
6684:
6683:
6681:
6677:
6669:
6668:
6664:
6663:
6662:
6659:
6657:
6654:
6652:
6649:
6647:
6644:
6642:
6639:
6635:
6632:
6631:
6630:
6629:Jacques Monod
6626:
6623:
6621:
6618:
6616:
6613:
6612:
6610:
6606:
6596:
6593:
6591:
6588:
6586:
6583:
6581:
6578:
6576:
6573:
6571:
6568:
6566:
6563:
6559:
6556:
6555:
6554:
6551:
6549:
6546:
6544:
6543:Homeotic gene
6541:
6540:
6538:
6534:
6528:
6525:
6523:
6520:
6518:
6515:
6513:
6510:
6508:
6505:
6503:
6500:
6498:
6495:
6494:
6492:
6488:
6485:
6481:
6475:
6472:
6470:
6467:
6465:
6462:
6460:
6457:
6455:
6452:
6451:
6449:
6445:
6439:
6436:
6434:
6431:
6429:
6426:
6422:
6419:
6417:
6414:
6413:
6412:
6411:Morphogenesis
6409:
6407:
6404:
6403:
6401:
6397:
6391:
6388:
6386:
6383:
6381:
6378:
6376:
6373:
6371:
6368:
6367:
6365:
6361:
6355:
6352:
6350:
6347:
6345:
6342:
6340:
6337:
6335:
6332:
6330:
6326:
6323:
6321:
6318:
6316:
6313:
6311:
6308:
6306:
6303:
6302:
6300:
6298:
6294:
6288:
6285:
6281:
6278:
6277:
6276:
6273:
6271:
6268:
6266:
6263:
6261:
6258:
6256:
6253:
6251:
6248:
6246:
6245:Reaction norm
6243:
6241:
6238:
6237:
6235:
6231:
6227:
6223:
6215:
6210:
6208:
6203:
6201:
6196:
6195:
6192:
6180:
6177:
6176:
6173:
6167:
6164:
6162:
6159:
6157:
6154:
6152:
6149:
6147:
6143:
6140:
6138:
6135:
6133:
6130:
6128:
6125:
6123:
6120:
6118:
6115:
6114:
6112:
6108:
6102:
6101:Sewall Wright
6099:
6097:
6094:
6092:
6089:
6088:
6086:
6082:
6076:
6073:
6071:
6068:
6066:
6063:
6061:
6058:
6056:
6053:
6052:
6050:
6048:
6047:Genetic drift
6044:
6038:
6035:
6033:
6030:
6029:
6027:
6019:
6013:
6010:
6008:
6005:
6003:
6000:
5998:
5995:
5994:
5992:
5990:
5986:
5980:
5977:
5975:
5972:
5970:
5967:
5965:
5962:
5960:
5957:
5955:
5952:
5950:
5947:
5945:
5942:
5940:
5937:
5935:
5932:
5930:
5927:
5925:
5922:
5920:
5917:
5915:
5912:
5910:
5907:
5906:
5904:
5900:
5896:
5889:
5884:
5882:
5877:
5875:
5870:
5869:
5866:
5854:
5851:
5849:
5846:
5844:
5841:
5839:
5836:
5835:
5833:
5829:
5823:
5820:
5818:
5815:
5813:
5810:
5808:
5805:
5803:
5800:
5799:
5797:
5793:
5789:
5785:
5778:
5773:
5771:
5766:
5764:
5759:
5758:
5755:
5749:
5745:
5742:
5739:
5737:
5734:
5733:
5729:
5722:
5718:
5714:
5710:
5706:
5702:
5698:
5694:
5689:
5685:
5679:
5675:
5670:
5669:
5664:
5645:
5641:
5635:
5628:
5627:
5619:
5616:
5611:
5607:
5603:
5599:
5595:
5591:
5584:
5581:
5576:
5572:
5568:
5564:
5560:
5556:
5549:
5547:
5543:
5537:
5534:
5521:
5517:
5513:
5509:
5505:
5501:
5497:
5493:
5489:
5485:
5481:
5477:
5470:
5467:
5463:
5459:
5455:
5450:
5445:
5441:
5437:
5433:
5429:
5425:
5418:
5415:
5410:
5406:
5402:
5398:
5394:
5390:
5383:
5380:
5367:
5363:
5362:
5361:The Telegraph
5357:
5350:
5347:
5335:
5331:
5325:
5321:
5317:
5316:
5308:
5305:
5293:
5289:
5283:
5279:
5275:
5274:
5269:
5262:
5259:
5254:
5250:
5245:
5240:
5236:
5232:
5228:
5225:(June 2006).
5224:
5218:
5215:
5210:
5204:
5200:
5199:
5191:
5188:
5183:
5177:
5173:
5172:
5164:
5161:
5156:
5149:
5145:
5140:
5135:
5130:
5125:
5121:
5117:
5116:PLOS Genetics
5113:
5106:
5103:
5098:
5094:
5089:
5084:
5079:
5074:
5070:
5066:
5062:
5058:
5054:
5047:
5044:
5029:
5025:
5021:
5017:
5013:
5009:
5005:
5001:
4997:
4990:
4983:
4980:
4975:
4971:
4966:
4961:
4957:
4953:
4949:
4945:
4941:
4934:
4931:
4926:
4920:
4916:
4912:
4911:
4903:
4900:
4895:
4891:
4887:
4883:
4879:
4875:
4872:(5): 467–73.
4871:
4867:
4860:
4857:
4852:
4848:
4844:
4840:
4837:(6): 353–72.
4836:
4832:
4825:
4822:
4817:
4813:
4808:
4803:
4799:
4795:
4791:
4787:
4783:
4776:
4773:
4768:
4764:
4759:
4754:
4749:
4744:
4740:
4736:
4733:(4): 1193–8.
4732:
4728:
4724:
4717:
4714:
4702:
4698:
4694:
4687:
4684:
4679:
4675:
4671:
4664:
4662:
4660:
4656:
4640:
4636:
4632:
4628:
4624:
4620:
4616:
4612:
4608:
4601:
4594:
4591:
4584:
4578:
4574:
4568:
4565:
4564:
4560:
4555:
4551:
4547:
4546:
4541:
4534:
4531:
4526:
4522:
4518:
4514:
4510:
4506:
4503:(2): 99–128.
4502:
4498:
4491:
4488:
4483:
4479:
4474:
4469:
4465:
4461:
4457:
4453:
4449:
4442:
4439:
4424:
4420:
4416:
4412:
4408:
4404:
4400:
4397:(4): 255–66.
4396:
4392:
4385:
4381:
4378:Visscher PM,
4374:
4372:
4370:
4366:
4361:
4357:
4353:
4349:
4345:
4341:
4337:
4333:
4326:
4323:
4308:
4304:
4300:
4296:
4292:
4288:
4284:
4277:
4270:
4267:
4255:
4251:
4247:
4243:
4239:
4234:
4233:2027.42/42804
4229:
4225:
4221:
4217:
4213:
4209:
4202:
4199:
4184:
4180:
4176:
4172:
4168:
4163:
4158:
4154:
4150:
4146:
4142:
4135:
4128:
4126:
4122:
4107:
4103:
4099:
4095:
4091:
4087:
4083:
4079:
4075:
4068:
4061:
4059:
4055:
4050:
4046:
4041:
4036:
4032:
4028:
4024:
4020:
4016:
4009:
4006:
4001:
3995:
3987:
3983:
3979:
3973:
3969:
3962:
3959:
3946:
3942:
3938:
3934:
3927:
3924:
3917:
3913:
3910:
3908:
3905:
3903:
3900:
3899:
3895:
3893:
3891:
3887:
3885:
3880:
3876:
3872:
3867:
3865:
3861:
3856:
3852:
3848:
3844:
3840:
3836:
3832:
3828:
3817:
3807:
3803:
3797:
3795:
3790:This section
3788:
3779:
3778:
3773:Controversies
3772:
3770:
3768:
3749:
3731:
3724:
3701:
3698:
3694:
3676:
3671:
3667:
3663:
3660:
3653:
3652:
3651:
3649:
3628:
3624:
3612:
3604:
3599:
3592:
3590:
3588:
3579:
3577:
3575:
3571:
3567:
3563:
3558:
3556:
3553:
3549:
3546:
3543:
3540:
3536:
3532:
3528:
3525:
3521:
3517:
3512:
3510:
3506:
3497:
3477:
3474:
3465:
3449:
3446:
3437:
3434:
3433:
3418:
3411:
3395:
3392:
3383:
3380:
3379:
3362:
3359:
3350:
3334:
3331:
3322:
3319:
3318:
3303:
3296:
3280:
3277:
3268:
3265:
3264:
3249:
3242:
3226:
3223:
3214:
3211:
3210:
3195:
3188:
3174:
3167:
3164:
3163:
3148:
3141:
3127:
3120:
3118:Relationship
3117:
3116:
3110:
3107:
3093:
3090:
3082:
3068:
3065:
3057:
3056:are found as
3043:
3023:
3000:
2995:
2991:
2987:
2984:
2979:
2975:
2971:
2968:
2960:
2959:
2958:
2956:
2938:
2934:
2911:
2907:
2897:
2883:
2861:
2858:
2854:
2831:
2827:
2804:
2800:
2791:
2774:
2771:
2768:
2763:
2760:
2756:
2752:
2747:
2743:
2739:
2734:
2730:
2726:
2723:
2720:
2715:
2712:
2708:
2700:
2699:
2698:
2692:
2690:
2688:
2684:
2679:
2662:
2656:
2653:
2650:
2644:
2641:
2633:
2630:
2627:
2621:
2615:
2607:
2603:
2599:
2594:
2590:
2583:
2579:
2573:
2568:
2564:
2555:
2537:
2533:
2527:
2524:
2495:
2491:
2487:
2482:
2478:
2472:
2469:
2460:
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6665:
6558:eyeless gene
6454:Evolvability
6428:Segmentation
6305:Canalisation
6275:Heterochrony
6265:Heritability
6264:
6233:Key concepts
6117:Biogeography
6091:R. A. Fisher
5969:Heritability
5968:
5902:Key concepts
5802:Heritability
5801:
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3890:heritability
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3871:twin studies
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103:twin studies
87:
81:, including
76:
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39:Heritability
38:
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6817:Tan Jiazhen
6812:Hans Stubbe
6749:Lysenkoists
6656:Mike Levine
6565:Distal-less
6390:Polyphenism
6370:Epigenetics
6222:development
6070:Coalescence
5268:"Chapter 5"
4338:(1): 1–27.
4162:10616/47311
3855:David Shenk
3851:David Moore
3827:Steven Rose
3814:August 2016
1758:Basic model
1714:correlation
1683:familiarity
1411:statistical
1405:Since only
1384:Assumptions
558:interaction
189:Phenotype (
32:differences
6919:Categories
6858:Lamarckism
6785:Dissidents
6742:Lysenkoism
6634:Lac operon
6459:Robustness
6438:Modularity
6433:Metamerism
6339:Plasticity
6334:Pleiotropy
6287:Heterotopy
6012:Ecological
6002:Artificial
5653:2016-04-02
5340:2016-04-02
5298:2016-04-02
5037:2020-09-06
4792:: 422–39.
4707:2020-02-20
4648:29 October
4552:(4): 382.
4432:2015-08-28
4316:2019-12-05
4260:2020-12-24
4192:2021-02-08
4115:2020-09-06
3918:References
3831:Jay Joseph
3794:unbalanced
3518:(REML) or
2166:sires and
1974:error term
1746:=1-r(MZ).
1695:Twin study
1658:regression
1647:Figure 2.
1527:twin study
1465:J. L. Lush
221:) + 2 Cov(
177:Definition
167:antibiotic
155:hair color
151:univariate
124:inbreeding
63:population
6585:Morphogen
6570:Engrailed
6553:Pax genes
6474:Tinkering
6320:Epistasis
6315:Dominance
6226:phenotype
6122:Evolution
5989:Selection
5807:Dominance
5713:0890-2070
5024:143217984
5016:1469-1825
4678:422371269
4627:0963-7214
4567:review of
4525:204981536
4497:Cognition
4303:1878-7304
4179:220260262
3994:cite book
3986:908409996
3943:(1): 29.
3806:talk page
3544:, WOMBAT
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3091:θ
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608:μ
542:), where
491:selection
364:epistatic
159:eye color
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6838:Heredity
6831:Concepts
6776:VASKhNIL
6548:Hox gene
6536:Elements
6517:Homeobox
6146:genomics
6084:Founders
5853:Heredity
5843:Genomics
5784:Genetics
5744:Archived
5721:41842465
5644:Archived
5610:26413946
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5409:27906501
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4212:Genetica
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4106:Archived
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3907:Heredity
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3864:Lewontin
3524:raw data
2095:′
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1457:C. C. Li
560:between
360:dominant
217:) + Var(
213:) = Var(
131:canalize
109:Overview
51:genetics
47:breeding
21:heredity
6679:Debates
6490:Systems
6416:Eyespot
6280:Neoteny
5997:Natural
5964:Fitness
5516:6797128
5508:1198102
5488:Bibcode
5480:Science
5449:5812976
5372:4 April
5223:Rose SP
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5065:Bibcode
4965:2226023
4915:Longman
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3605:=R/S.
2252:d.f.
1963:error
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1423:twins
383:locus
119:genes
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6144:and
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5634:ISBN
5606:PMID
5528:2021
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5405:PMID
5374:2018
5324:ISBN
5282:ISBN
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4674:OCLC
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4348:PMID
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4090:PMID
4045:PMID
4000:link
3982:OCLC
3972:ISBN
3953:2015
3862:and
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3250:0
3227:2
3224:1
3196:1
3175:1
3128:r
3094:=
3069:=
3066:r
3024:r
3001:,
2996:d
2992:V
2985:+
2980:a
2976:V
2972:r
2969:=
2939:d
2935:V
2912:a
2908:V
2884:e
2862:j
2859:i
2855:d
2832:j
2805:i
2775:,
2772:e
2769:+
2764:j
2761:i
2757:d
2753:+
2748:j
2740:+
2735:i
2727:+
2721:=
2716:j
2713:i
2709:y
2663:W
2660:)
2657:1
2651:r
2648:(
2645:+
2642:S
2637:)
2634:W
2628:S
2625:(
2622:4
2616:=
2608:e
2604:V
2600:+
2595:g
2591:V
2584:g
2580:V
2574:=
2569:2
2565:H
2538:g
2534:V
2528:4
2525:1
2496:e
2492:V
2488:+
2483:g
2479:V
2473:4
2470:3
2447:W
2426:)
2423:1
2417:r
2414:(
2411:n
2385:)
2379:g
2375:V
2369:4
2366:1
2360:(
2357:r
2354:+
2349:e
2345:V
2341:+
2336:g
2332:V
2326:4
2323:3
2300:S
2279:1
2273:n
2226:e
2222:V
2199:g
2195:V
2174:r
2154:n
2134:,
2120:g
2116:V
2110:4
2107:1
2102:=
2099:)
2092:e
2088:+
2085:g
2080:2
2077:1
2072:+
2066:,
2063:e
2060:+
2057:g
2052:2
2049:1
2044:+
2038:(
2034:r
2031:r
2028:o
2025:c
2021:=
2018:)
2011:z
2007:,
2004:z
2001:(
1997:r
1994:r
1991:o
1988:c
1937:e
1934:+
1929:i
1925:g
1919:2
1916:1
1911:+
1905:=
1900:i
1896:z
1869:e
1858:i
1856:G
1840:i
1836:g
1812:e
1809:+
1804:i
1800:g
1796:+
1790:=
1785:i
1781:y
1766:i
1764:G
1744:e
1740:e
1736:h
1732:c
1728:c
1721:H
1633:t
1629:b
1621:r
1602:r
1599:t
1594:=
1589:r
1586:b
1581:=
1576:2
1572:h
1467:(
1459:(
1407:P
1389:"
1359:=
1354:B
1351:B
1347:d
1343:)
1340:B
1337:B
1334:(
1331:f
1328:+
1323:b
1320:B
1316:d
1312:)
1309:b
1306:B
1303:(
1300:f
1297:+
1292:b
1289:b
1285:d
1281:)
1278:b
1275:b
1272:(
1269:f
1246:,
1241:2
1236:B
1233:B
1229:d
1225:)
1222:B
1219:B
1216:(
1213:f
1210:+
1205:2
1200:b
1197:B
1193:d
1189:)
1186:b
1183:B
1180:(
1177:f
1174:+
1169:2
1164:b
1161:b
1157:d
1153:)
1150:b
1147:b
1144:(
1141:f
1138:=
1135:)
1132:D
1129:(
1125:r
1122:a
1119:V
1090:=
1085:B
1082:B
1078:a
1074:)
1071:B
1068:B
1065:(
1062:f
1059:+
1054:b
1051:B
1047:a
1043:)
1040:b
1037:B
1034:(
1031:f
1028:+
1023:b
1020:b
1016:a
1012:)
1009:b
1006:b
1003:(
1000:f
977:,
972:2
967:B
964:B
960:a
956:)
953:B
950:B
947:(
944:f
941:+
936:2
931:b
928:B
924:a
920:)
917:b
914:B
911:(
908:f
905:+
900:2
895:b
892:b
888:a
884:)
881:b
878:b
875:(
872:f
869:=
866:)
863:A
860:(
856:r
853:a
850:V
816:.
813:)
810:)
805:j
801:B
795:i
791:B
787:(
781:=
776:j
773:i
769:d
765:(
757:+
754:)
751:)
746:j
742:B
738:+
733:i
729:B
725:(
719:=
714:j
711:i
707:a
703:(
695:+
687:=
677:)
672:j
668:B
662:i
658:B
654:(
647:+
644:)
639:j
635:B
631:+
626:i
622:B
618:(
611:+
605:=
596:j
593:i
589:P
572:j
569:B
565:i
562:B
554:j
551:B
547:i
544:B
540:j
537:B
535:,
533:i
530:B
526:B
476:h
472:H
452:)
449:P
446:(
442:r
439:a
436:V
430:)
427:A
424:(
420:r
417:a
414:V
407:=
402:2
398:h
356:H
337:)
334:P
331:(
327:r
324:a
321:V
315:)
312:G
309:(
305:r
302:a
299:V
292:=
287:2
283:H
259:,
254:2
250:H
239:E
237:,
235:G
227:E
225:,
223:G
219:E
215:G
211:P
199:E
195:G
191:P
23:.
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