133:
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,
4116:
38:
5166:
128:. 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
525:
116:). 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.
5473:...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'.
3609:
3899:. 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,
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841:
1655:
3860:, 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,
504:. 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
3514:
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
590:
3595:
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
1535:
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
396:
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
175:
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
3707:
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
1666:
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
1718:
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
1696:– 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.
132:
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
3868:
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
3541:
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
99:
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
3513:
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
3703:
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.
2143:
3708:
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.
96:. 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.
1404:" 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
836:{\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}}}
1711:
1741:, 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,
1691:
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
3522:) 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.
1443:
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
492:
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
3537:
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.
137:, 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.
3600:. 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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998:
1399:
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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159:
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
164:– 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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476:
361:
80:
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
2406:
5376:
2796:
3542:
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.
1895:
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
1723:
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
1383:
1114:
3774:
1625:
595:
1993:
1958:
5654:
1432:
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
2689:. 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.
2519:
144:
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
1555:. 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.
3022:
567:
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
1420:
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
2561:
1833:
5939:
3501:
3473:
3419:
3386:
3358:
3304:
3250:
4144:
3856:. Bentall has claimed that such heritability scores are typically calculated counterintuitively to derive numerically high scores, that heritability is misinterpreted as
1773:
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
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2828:
388:
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
3698:
3115:
2910:
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
152:
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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3170:
3065:
2885:
2447:
280:
5785:
2962:
2935:
2249:
2222:
1863:
4010:
2300:
3090:
5896:
3440:
3325:
3271:
3217:
3196:
3149:
3045:
2905:
2468:
2321:
2195:
2175:
1890:
3525:
When a large, complex pedigree or another aforementioned type of data is available, heritability and other quantitative genetic parameters can be estimated by
5062:
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).
1540:
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
140:
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
1972:). 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
3955:
3877:
emphasize that heritability is itself a function of environmental variation. However, some researchers argue that it is possible to disentangle the two.
1513:
of breeding studies, using the intraclass correlation of relatives. Various methods of estimating components of variance (and, hence, heritability) from
1124:
855:
1968:
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
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to estimate the influence on a trait by genetic factors, which is reflected by the rate and influence of putatively associated genetic loci (usually
4840:
Cattell RB (November 1960). "The multiple abstract variance analysis equations and solutions: for nature-nurture research on continuous variables".
4951:"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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1714:
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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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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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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5692:
5648:
5338:
5296:
5217:
5190:
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1662:'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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6743:
403:
288:
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5302:
6677:
5778:
3711:
Observing the response to selection in an artificial selection experiment will allow calculation of realized heritability as in Fig. 4.
2327:
6250:
369:
is the broad-sense heritability. This reflects all the genetic contributions to a population's phenotypic variance including additive,
6522:
4933:
4649:
3885:
1984:
1525:
6222:
5889:
6265:
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172:. Environment and genetics may also interact, and heritability analyses can test for and examine these interactions (GxE models).
6479:
5984:
3519:
3515:
1521:
1506:
148:, which makes heritability difficult to measure in some cases. Recent insights in molecular biology have identified changes in
4264:
45:
in height between people. This is not the same as asking to what extent do genetic factors influence height in any one person.
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5771:
5754:
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1680:
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1006:
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6507:
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3526:
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3717:
3559:
2138:{\displaystyle \mathrm {corr} (z,z')=\mathrm {corr} (\mu +{\frac {1}{2}}g+e,\mu +{\frac {1}{2}}g+e')={\frac {1}{4}}V_{g}}
1577:
6945:
5882:
3893:
3556:
1552:
6474:
5964:
4793:"Nurture net of nature: Re-evaluating the role of shared environments in academic achievement and verbal intelligence"
1901:
1635:
1437:
1436:, siblings, parents and offspring, rather than from more distantly related (and therefore less similar) subjects. The
1448:
is generally not possible when gathering human data, relying on naturally occurring relationships and environments.
532:
The simplest genetic model involves a single locus with two alleles (b and B) affecting one quantitative phenotype.
6666:
6532:
6364:
1672:
1483:
37:
6085:
4341:
Maccoby EE (February 2000). "Parenting and its effects on children: on reading and misreading behavior genetics".
1551:, or because of evocative effects (where a genome evokes environments by its effect on them), G and E may covary:
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6359:
5959:
5832:
5371:
5330:
1429:
89:
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contributions of genes and environment, as such traits result from multiple causes interacting. In particular,
528:
Figure 1. Relationship of phenotypic values to additive and dominance effects using a completely dominant locus.
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6137:
5822:
3943:
2474:
149:
1765:
The second set of methods of estimation of heritability involves ANOVA and estimation of variance components.
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6600:
6537:
6512:
6395:
6315:
5949:
3553:
1498:
1451:
In classical quantitative genetics, there were two schools of thought regarding estimation of heritability.
141:
5165:
2975:
6585:
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6339:
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3624:
of plants and animals, the expected response to selection of a trait with known narrow-sense heritability
3597:
3580:
2564:
370:
4459:"The paradox of intelligence: Heritability and malleability coexist in hidden gene-environment interplay"
4394:
3612:
Figure 4. Strength of selection (S) and response to selection (R) in an artificial selection experiment,
2529:
1719:
heritability". Fraternal or dizygotic (DZ) twins on average share half their genes (assuming there is no
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6696:
6590:
6517:
6416:
6354:
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6280:
6215:
6176:
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5798:
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1786:
1502:
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1471:
1463:
145:
101:
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Gielen M, Lindsey PJ, Derom C, Smeets HJ, Souren NY, Paulussen AD, Derom R, Nijhuis JG (January 2008).
1728:
5752:
Quantitative
Genetics Resources website, including the two volume book by Lynch and Walsh. Free access
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1401:
513:
156:
5702:
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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93:
5064:"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
524:
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105:
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5103:
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4857:
4822:
4773:
4684:
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4417:
4358:
4309:
4248:
4177:
4100:
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3992:
3982:
3816:
3667:
3584:
3572:
3097:
2697:
1475:
1388:
501:
193:
184:. In practice, all human behavioral traits vary and almost all traits show some heritability.
125:
109:
77:
3627:
3155:
3050:
6858:
6661:
6630:
6625:
6580:
6208:
6106:
6017:
5711:
5608:
5573:
5506:
5454:
5446:
5407:
5249:
5180:
5144:
5134:
5093:
5083:
5014:
4970:
4962:
4884:
4849:
4812:
4804:
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4753:
4625:
4564:
4515:
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4409:
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4301:
4238:
4230:
4167:
4159:
4092:
4045:
4037:
3576:
2860:
2417:
393:
255:
216:
Likewise the phenotypic variance in the trait – Var (P) – is the sum of effects as follows:
69:
57:
4610:
3804:
2940:
2913:
2227:
2200:
1841:
6832:
6812:
6671:
6575:
6385:
5924:
5758:
4390:
4145:"High-definition likelihood inference of genetic correlations across human complex traits"
3853:
3845:
2965:
1979:
The second group of progeny are comparisons of means of half sibs with each other (called
494:
378:
41:
Studies of heritability ask questions such as to what extent do genetic factors influence
2279:
1563:
The first school of estimation uses regression and correlation to estimate heritability.
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Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
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4734:"The mystery of missing heritability: Genetic interactions create phantom heritability"
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The use of ANOVA to calculate heritability often fails to account for the presence of
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Heritability measures the fraction of phenotype variability that can be attributed to
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6255:
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6101:
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DeFries JC, Fulker DW (September 1985). "Multiple regression analysis of twin data".
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381:, where individuals are directly affected by their parents' phenotype, such as with
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1262:{\displaystyle \mathrm {Var} (D)=f(bb)d_{bb}^{2}+f(Bb)d_{Bb}^{2}+f(BB)d_{BB}^{2},}
993:{\displaystyle \mathrm {Var} (A)=f(bb)a_{bb}^{2}+f(Bb)a_{Bb}^{2}+f(BB)a_{BB}^{2},}
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5139:
4354:
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Hill WG, Goddard ME, Visscher PM (February 2008). MacKay TF, Goddard ME (eds.).
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1724:
1710:
1679:, the slope is always less than one). This regression effect also underlies the
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5326:
Doctoring the Mind: Is Our
Current Treatment of Mental Illness Really Any Good?
5068:
Proceedings of the
National Academy of Sciences of the United States of America
5000:"Insensitivity of the analysis of variance to heredity-environment interaction"
4738:
Proceedings of the
National Academy of Sciences of the United States of America
4703:
4305:
4041:
3566:
1753:, reflects the degree to which identical twins raised together are dissimilar,
1531:
Studies of human heritability often utilize adoption study designs, often with
17:
6868:
6752:
6644:
6344:
6297:
5746:
5123:"Data and theory point to mainly additive genetic variance for complex traits"
5018:
4966:
4163:
3841:
1705:
1537:
1482:). It is based on the analysis of correlations and, by extension, regression.
1421:
177:
165:
161:
134:
113:
73:
5723:
5026:
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4637:
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3996:
2682:{\displaystyle H^{2}={\frac {V_{g}}{V_{g}+V_{e}}}={\frac {4(S-W)}{S+(r-1)W}}}
6595:
6563:
6330:
6236:
6132:
5510:
5088:
4758:
4629:
4569:
4550:
374:
169:
53:
27:
Estimation of effect of genetic variation on phenotypic variation of a trait
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5468:
5450:
5419:
5263:
5158:
5107:
4984:
4861:
4826:
4777:
4492:
4421:
4362:
4181:
4104:
4059:
5518:
4896:
4527:
4252:
3880:
The controversy over heritability estimates is largely via their basis in
1118:
There is a similar relationship for the variance of dominance deviations:
6848:
6786:
6558:
6527:
5863:
5853:
5794:
5254:
5237:
4243:
3917:
3534:
1467:
1425:
61:
31:
6290:
4925:
4888:
4474:
4234:
4172:
3562:
3120:
Some common relationships and their coefficients are given in Table 2.
5874:
5747:
Stanford
Encyclopedia of Philosophy entry on Heredity and Heritability
5435:"Missing compared to what? Revisiting heritability, genes and culture"
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5411:
5288:
4853:
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389:
181:
5715:
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2964:
as the dominance deviation variance, intraclass correlations become
471:{\displaystyle h^{2}={\frac {\mathrm {Var} (A)}{\mathrm {Var} (P)}}}
356:{\displaystyle H^{2}={\frac {\mathrm {Var} (G)}{\mathrm {Var} (P)}}}
4078:"Concepts, estimation and interpretation of SNP-based heritability"
3545:
The pedigrees can be viewed using programs such as Pedigree Viewer
88:
Other causes of measured variation in a trait are characterized as
5238:"Commentary: heritability estimates--long past their sell-by date"
3607:
2401:{\displaystyle {\frac {3}{4}}V_{g}+V_{e}+r({{\frac {1}{4}}V_{g}})}
1653:
1520:
Today, heritability can be estimated from general pedigrees using
1433:
36:
180:, or because they are omni-present, like if everyone is drinking
6568:
4076:
Yang J, Zeng J, Goddard ME, Wray NR, Visscher PM (August 2017).
1541:
1532:
1440:
for heritability estimates is improved with large sample sizes.
382:
176:
in the population, such as no one having access to a particular
129:
6725:
6204:
5878:
5767:
5551:
Tredoux, Gavan. "The Nature and Nurture of Rectangles." (2019).
4506:
Block N (August 1995). "How heritability misleads about race".
4395:"Heritability in the genomics era--concepts and misconceptions"
1544:, potentially resulting in an underestimation of heritability.
252:) can be controlled and held at 0. In this case, heritability,
5398:
Moore DS, Shenk D (January 2017). "The heritability fallacy".
3787:
2197:
progeny per sire, we can calculate the following ANOVA, using
508:
leads to an estimate of the narrow-sense heritability (called
497:
in which heritability can be estimated and selection modeled.
6721:
4586:. Sunderland, MA: Sinauer and Associates. pp. xv + 652.
3571:
Pedigree models are helpful for untangling confounds such as
2791:{\displaystyle y_{ij}=\mu +\alpha _{i}+\alpha _{j}+d_{ij}+e,}
2700:
for testing for interaction effects than for direct effects.
1683:
for analyzing twins selected for one member being affected.
2149:
since environmental effects are independent of each other.
3124:
Table 2: Coefficients for calculating variance components
3977:
Gazzaniga MS, Heatherton TF, Halpern DF (February 2015).
3714:
Heritability in the above equation is equal to the ratio
1571:
In the comparison of relatives, we find that in general,
4732:
Zuk O, Hechter E, Sunyaev SR, Lander ES (January 2012).
4024:
Yang J, Lee SH, Goddard ME, Visscher PM (January 2011).
3776:
only if the genotype and the environmental noise follow
3812:
5179:
Plomin R, DeFries JC, McClearn GE, McGuffin P (2017).
1378:{\displaystyle f(bb)d_{bb}+f(Bb)d_{Bb}+f(BB)d_{BB}=0.}
1109:{\displaystyle f(bb)a_{bb}+f(Bb)a_{Bb}+f(BB)a_{BB}=0.}
4026:"GCTA: a tool for genome-wide complex trait analysis"
3720:
3670:
3630:
3482:
3454:
3428:
3400:
3367:
3339:
3313:
3285:
3259:
3231:
3205:
3184:
3158:
3137:
3100:
3075:
3053:
3033:
2978:
2943:
2916:
2893:
2863:
2836:
2809:
2717:
2573:
2532:
2477:
2456:
2420:
2330:
2309:
2282:
2230:
2203:
2183:
2163:
1996:
1904:
1878:
1844:
1789:
1580:
1278:
1127:
1009:
858:
593:
406:
291:
258:
4683:(1st ed.). Ames, Iowa: Iowa State Univ. Press.
4611:"Three Laws of Behavior Genetics and What They Mean"
2887:
is the dominance deviation for the ij genotype, and
6887:
6841:
6795:
6759:
6689:
6618:
6546:
6500:
6493:
6457:
6409:
6373:
6306:
6243:
6120:
6094:
6056:
6031:
5998:
5912:
5841:
5805:
3836:Heritability estimates' prominent critics, such as
3769:{\displaystyle \mathrm {Var} (A)/\mathrm {Var} (P)}
1620:{\displaystyle h^{2}={\frac {b}{r}}={\frac {t}{r}}}
845:The additive genetic variance at this locus is the
5400:Wiley Interdisciplinary Reviews: Cognitive Science
4674:
4672:
4670:
4384:
4382:
4380:
3768:
3692:
3649:
3495:
3467:
3434:
3413:
3380:
3352:
3319:
3298:
3265:
3244:
3211:
3190:
3164:
3143:
3109:
3084:
3059:
3039:
3016:
2956:
2929:
2899:
2879:
2849:
2822:
2790:
2681:
2555:
2513:
2462:
2441:
2400:
2315:
2294:
2243:
2216:
2189:
2169:
2137:
1952:
1884:
1857:
1827:
1619:
1377:
1261:
1108:
992:
835:
470:
355:
274:
5559:
5557:
3587:, shared environment, and maternal gene effects.
196:as the sum of genetic and environmental effects:
4551:"Principles of Population Genetics, 4th edition"
1953:{\displaystyle z_{i}=\mu +{\frac {1}{2}}g_{i}+e}
85:explained by the environment or random chance?"
5485:Marcus W. Feldman; Richard C. Lewontin (1975).
1509:, as well as other schools. It is based on the
1391:of phenotype on genotype is shown in Figure 1.
392:. Since each parent passes a single allele per
539:alleles can be 0, 1, or 2. For any genotype, (
485:is used to denote broad sense, and lower case
6737:
6216:
5890:
5779:
3848:, focus largely on heritability estimates in
1671:," since the offspring values always tend to
8:
5685:Genetics and analysis of quantitative traits
1667:the slope. (This is the source of the term "
1559:Regression/correlation methods of estimation
5599:Turkheimer E (2015). "Genetic Prediction".
4618:Current Directions in Psychological Science
2567:between half sibs. We can easily calculate
6744:
6730:
6722:
6497:
6223:
6209:
6201:
5897:
5883:
5875:
5786:
5772:
5764:
4009:: CS1 maint: location missing publisher (
1761:Analysis of variance methods of estimation
5458:
5433:Feldman MW, Ramachandran S (April 2018).
5253:
5148:
5138:
5097:
5087:
4974:
4816:
4767:
4757:
4568:
4482:
4242:
4171:
4049:
3746:
3741:
3721:
3719:
3681:
3669:
3638:
3629:
3483:
3481:
3455:
3453:
3427:
3401:
3399:
3368:
3366:
3340:
3338:
3312:
3286:
3284:
3258:
3232:
3230:
3204:
3183:
3157:
3136:
3099:
3074:
3052:
3032:
3005:
2989:
2977:
2948:
2942:
2921:
2915:
2892:
2868:
2862:
2841:
2835:
2814:
2808:
2770:
2757:
2744:
2722:
2716:
2629:
2617:
2604:
2593:
2587:
2578:
2572:
2547:
2533:
2531:
2514:{\displaystyle {\frac {3}{4}}V_{g}+V_{e}}
2505:
2492:
2478:
2476:
2455:
2419:
2388:
2374:
2373:
2358:
2345:
2331:
2329:
2308:
2281:
2235:
2229:
2208:
2202:
2182:
2162:
2129:
2115:
2085:
2057:
2034:
1997:
1995:
1938:
1924:
1909:
1903:
1877:
1849:
1843:
1813:
1794:
1788:
1607:
1594:
1585:
1579:
1360:
1329:
1298:
1277:
1250:
1242:
1214:
1206:
1178:
1170:
1128:
1126:
1091:
1060:
1029:
1008:
981:
973:
945:
937:
909:
901:
859:
857:
814:
804:
782:
770:
755:
742:
720:
708:
700:
681:
671:
663:
648:
635:
627:
602:
594:
592:
445:
423:
420:
411:
405:
330:
308:
305:
296:
290:
263:
257:
5185:(2nd ed.). New York: W.H. Freeman.
4918:Falconer DS, Mackay TF (December 1995).
3122:
2857:is the additive effect of the j allele,
2830:is the additive effect of the i allele,
2253:
1709:
849:of the squares of the additive effects:
523:
6910:Suppressed research in the Soviet Union
6864:Inheritance of acquired characteristics
4143:Ning Z, Pawitan Y, Shen X (June 2020).
3934:
2704:Model with additive and dominance terms
1493:The second was originally developed by
377:(multi-genic interactions), as well as
6713:Index of evolutionary biology articles
6190:Index of evolutionary biology articles
5564:Turkheimer E (2011). "Still missing".
5379:from the original on 28 September 2011
4002:
5242:International Journal of Epidemiology
5209:Introduction to quantitative genetics
4921:Introduction to Quantitative Genetics
4791:Daw J, Guo G, Harris KM (July 2015).
4681:An introduction to genetic statistics
3548:, and analyzed with programs such as
3017:{\displaystyle =rV_{a}+\theta V_{d},}
2937:as the additive genetic variance and
1642:is the coefficient of regression and
1536:second and more common design is the
1490:as a way of estimating heritability.
7:
6900:Collectivization in the Soviet Union
5687:. Sunderland, Mass.: Sinauer Assoc.
4655:from the original on 19 October 2013
4138:
4136:
4071:
4069:
3558:, MCMCglmm within the R environment
1542:not completely genetically identical
512:). This is the principle underlying
5643:. New York: Routledge. p. 81.
4708:Stanford Encyclopedia of Philosophy
4702:Stephen Downes and Lucas Matthews.
4457:Sauce B, Matzel LD (January 2018).
4219:"Coming to terms with heritability"
2556:{\displaystyle {\frac {1}{4}}V_{g}}
2255:Table 1: ANOVA for Sire experiment
1646:is the coefficient of correlation.
500:Additive variance is important for
6523:Evolutionary developmental biology
4287:"The intelligence of heritability"
4030:American Journal of Human Genetics
3958:from the original on 2 August 2015
3753:
3750:
3747:
3728:
3725:
3722:
2044:
2041:
2038:
2035:
2007:
2004:
2001:
1998:
1828:{\displaystyle y_{i}=\mu +g_{i}+e}
1737:The effect of shared environment,
1424:analyses required to estimate the
1135:
1132:
1129:
866:
863:
860:
452:
449:
446:
430:
427:
424:
337:
334:
331:
315:
312:
309:
25:
5806:Concepts in Quantitative Genetics
4584:Principles of Population Genetics
2968:of these parameters. In general,
2696:, because ANOVA has a much lower
1749:. Unique environmental variance,
6480:Evolution of sexual reproduction
5533:from the original on 20 May 2021
5212:(4th ed.). Essex: Longman.
5164:
4998:Wahlsten, Douglas (March 1990).
4809:10.1016/j.ssresearch.2015.02.011
3903:is a concept widely applicable.
3792:
1528:estimated from genetic markers.
192:Any particular phenotype can be
5704:European Journal of Personality
5660:from the original on 2016-04-04
5347:from the original on 2020-10-05
5305:from the original on 2017-07-19
5206:Falconer DS, Mackay TF (1998).
5044:from the original on 2020-10-05
4714:from the original on 2020-02-25
4439:from the original on 2016-03-24
4323:from the original on 2018-10-24
4267:from the original on 2020-12-02
4199:from the original on 2021-04-15
4122:from the original on 2020-10-05
3944:"Estimating Trait Heritability"
3520:single-nucleotide polymorphisms
3516:genome-wide association studies
3496:{\displaystyle {\frac {1}{16}}}
2251:as the environmental variance:
1507:North Carolina State University
397:heritability and is defined as
6251:Genotype–phenotype distinction
5990:Constructive neutral evolution
3763:
3757:
3738:
3732:
3644:
3631:
3468:{\displaystyle {\frac {1}{4}}}
3414:{\displaystyle {\frac {1}{8}}}
3381:{\displaystyle {\frac {1}{4}}}
3353:{\displaystyle {\frac {1}{2}}}
3299:{\displaystyle {\frac {1}{4}}}
3245:{\displaystyle {\frac {1}{2}}}
2694:gene–-environment interactions
2670:
2658:
2647:
2635:
2436:
2424:
2395:
2370:
2109:
2048:
2028:
2011:
1727:between MZ and DZ twins, i.e.
1353:
1344:
1322:
1313:
1291:
1282:
1235:
1226:
1199:
1190:
1163:
1154:
1145:
1139:
1084:
1075:
1053:
1044:
1022:
1013:
966:
957:
930:
921:
894:
885:
876:
870:
823:
820:
797:
775:
764:
761:
735:
713:
687:
664:
654:
628:
462:
456:
440:
434:
347:
341:
325:
319:
155:Estimates of heritability use
1:
6508:Regulation of gene expression
5566:Research in Human Development
5182:Behavioral Genetics: A Primer
5007:Behavioral and Brain Sciences
4609:Turkheimer E (October 2000).
3527:restricted maximum likelihood
1892:is the environmental effect.
1567:Comparison of close relatives
1466:, and further popularized by
379:maternal and paternal effects
64:that estimates the degree of
6678:Endless Forms Most Beautiful
6458:Evolution of genetic systems
6266:Gene–environment correlation
6261:Gene–environment interaction
5940:Fisher's fundamental theorem
5578:10.1080/15427609.2011.625321
5367:"Doctoring the Mind: Review"
5140:10.1371/journal.pgen.1000008
4520:10.1016/0010-0277(95)00678-r
4355:10.1146/annurev.psych.51.1.1
4217:Stoltenberg SF (June 1997).
3892:studies' conclusions is the
3888:studies to corroborate such
3815:or discuss the issue on the
2224:as the genetic variance and
1553:gene environment correlation
1517:are used in these analyses.
244:In a planned experiment Cov(
6657:Christiane NĂĽsslein-Volhard
5965:Coefficient of relationship
4582:Hartl DL, Clark AG (2007).
4343:Annual Review of Psychology
3942:Wray N, Visscher P (2008).
3657:can be estimated using the
2850:{\displaystyle \alpha _{j}}
2823:{\displaystyle \alpha _{i}}
1650:Parent-offspring regression
1499:The University of Edinburgh
6962:
6533:Hedgehog signaling pathway
6410:Developmental architecture
5601:The Hastings Center Report
5487:"The Heritability Hang–Up"
4306:10.1037/0708-5591.35.3.244
4042:10.1016/j.ajhg.2010.11.011
3981:(5th ed.). New York.
3807:towards certain viewpoints
1865:is the effect of genotype
1703:
1675:value for the population,
1636:coefficient of relatedness
29:
6915:Politicization of science
6710:
6360:Transgressive segregation
6185:
5960:Coefficient of inbreeding
5833:Effective population size
5683:Lynch M, Walsh B (1998).
5365:McGrath M (5 July 2009).
5331:New York University Press
5019:10.1017/S0140525X00077797
4967:10.1007/s10519-007-9170-3
4164:10.1038/s41588-020-0653-y
3884:. The scarce success of
1634:can be thought of as the
1464:The University of Chicago
772:Dominance Deviation
495:liability threshold model
6138:Evolutionary game theory
5920:Hardy–Weinberg principle
5823:Quantitative trait locus
4402:Nature Reviews. Genetics
4393:, Wray NR (April 2008).
3693:{\displaystyle R=h^{2}S}
3110:{\displaystyle \theta =}
30:Not to be confused with
6895:Bourgeois pseudoscience
6538:Notch signaling pathway
6513:Gene regulatory network
6396:Dual inheritance theory
5950:Shifting balance theory
5511:10.1126/science.1198102
5089:10.1073/pnas.1510497113
4797:Social Science Research
4759:10.1073/pnas.1119675109
4710:. Stanford University.
4630:10.1111/1467-8721.00084
3650:{\displaystyle (h^{2})}
3165:{\displaystyle \theta }
3060:{\displaystyle \theta }
1964:Intraclass correlations
1412:Estimating heritability
385:production in mammals.
6586:cis-regulatory element
6494:Control of development
6374:Non-genetic influences
6340:evolutionary landscape
5935:Linkage disequilibrium
5451:10.1098/rstb.2017.0064
4463:Psychological Bulletin
3778:Gaussian distributions
3770:
3694:
3651:
3617:
3598:linkage disequilibrium
3497:
3469:
3436:
3415:
3382:
3354:
3321:
3300:
3267:
3246:
3213:
3192:
3166:
3145:
3111:
3086:
3061:
3041:
3018:
2972:Intraclass correlation
2958:
2931:
2901:
2881:
2880:{\displaystyle d_{ij}}
2851:
2824:
2792:
2683:
2565:intraclass correlation
2557:
2515:
2464:
2443:
2442:{\displaystyle n(r-1)}
2402:
2317:
2296:
2245:
2218:
2191:
2171:
2157:In an experiment with
2139:
1983:). In addition to the
1954:
1886:
1859:
1829:
1715:
1663:
1621:
1379:
1263:
1110:
994:
837:
529:
472:
357:
276:
275:{\displaystyle H^{2},}
56:used in the fields of
46:
6941:Quantitative genetics
6874:Mendelian inheritance
6697:Nature versus nurture
6601:Cell surface receptor
6518:Evo-devo gene toolkit
6417:Developmental biology
6355:Polygenic inheritance
6281:Quantitative genetics
6177:Quantitative genetics
6086:Balding–Nichols model
6071:Population bottleneck
6066:Small population size
5970:Selection coefficient
5799:Quantitative genetics
4679:Kempthorne O (1957).
4570:10.1093/jhered/esm035
3979:Psychological science
3858:genetic determination
3771:
3695:
3652:
3611:
3604:Response to selection
3583:, and confounding of
3498:
3470:
3446:Double First Cousins
3437:
3416:
3383:
3355:
3322:
3301:
3268:
3247:
3214:
3193:
3167:
3146:
3112:
3087:
3062:
3042:
3019:
2959:
2957:{\displaystyle V_{d}}
2932:
2930:{\displaystyle V_{a}}
2902:
2882:
2852:
2825:
2793:
2684:
2558:
2516:
2465:
2444:
2403:
2318:
2297:
2269:Expected Mean Square
2246:
2244:{\displaystyle V_{e}}
2219:
2217:{\displaystyle V_{g}}
2192:
2172:
2140:
1955:
1887:
1860:
1858:{\displaystyle g_{i}}
1830:
1713:
1681:DeFries–Fulker method
1657:
1622:
1549:observational studies
1503:Iowa State University
1480:Iowa State University
1472:University of Chicago
1380:
1264:
1111:
995:
838:
710:Additive Effect
527:
510:realized heritability
506:response to selection
473:
358:
277:
146:phenotypic plasticity
102:quantitative genetics
100:important concept in
90:environmental factors
40:
6936:Genetic epidemiology
6606:Transcription factor
6321:Genetic assimilation
6308:Genetic architecture
6048:Background selection
6035:on genomic variation
6033:Effects of selection
5985:Population structure
5859:Evolutionary biology
5333:. pp. 123–127.
4842:Psychological Review
3895:missing heritability
3718:
3668:
3628:
3591:Genomic heritability
3581:prenatal environment
3480:
3452:
3426:
3398:
3365:
3337:
3311:
3283:
3257:
3229:
3203:
3182:
3156:
3135:
3098:
3073:
3051:
3031:
2976:
2941:
2914:
2907:is the environment.
2891:
2861:
2834:
2807:
2715:
2571:
2530:
2475:
2454:
2418:
2328:
2307:
2280:
2274:Between sire groups
2228:
2201:
2181:
2161:
1994:
1902:
1876:
1842:
1787:
1578:
1511:analysis of variance
1446:experimental control
1406:academic achievement
1402:missing heritability
1276:
1125:
1007:
856:
591:
514:artificial selection
404:
289:
256:
157:statistical analyses
6946:Population genetics
6808:Georgii Karpechenko
6702:Morphogenetic field
6619:Influential figures
6167:Population genomics
6043:Genetic hitchhiking
5930:Identity by descent
5906:Population genetics
5849:Population genetics
5503:1975Sci...190.1163F
5497:(4220): 1163–1168.
5323:Bentall RP (2009).
5080:2016PNAS..113.7377H
4750:2012PNAS..109.1193Z
4556:Journal of Heredity
4294:Canadian Psychology
4285:Wahlsten D (1994).
3913:Behavioral genetics
3850:behavioral sciences
3813:improve the article
3565:family of programs
3509:Linear mixed models
3125:
2412:Within sire groups
2295:{\displaystyle n-1}
2256:
1673:regress to the mean
1526:genomic relatedness
1522:linear mixed models
1255:
1219:
1183:
986:
950:
914:
94:observational error
6391:Genomic imprinting
6153:Landscape genetics
5757:2006-02-06 at the
5607:(5 Suppl): S32–8.
5445:(1743): 20170064.
5255:10.1093/ije/dyl064
4889:10.1007/BF01066239
4475:10.1037/bul0000131
4235:10.1007/BF02259512
3923:Heritability of IQ
3890:population-genetic
3766:
3690:
3659:breeder's equation
3647:
3622:selective breeding
3618:
3493:
3465:
3432:
3411:
3378:
3350:
3317:
3296:
3263:
3242:
3209:
3188:
3162:
3141:
3123:
3107:
3085:{\displaystyle r=}
3082:
3057:
3037:
3014:
2954:
2927:
2897:
2877:
2847:
2820:
2788:
2679:
2553:
2511:
2460:
2439:
2398:
2313:
2292:
2254:
2241:
2214:
2187:
2167:
2135:
1950:
1882:
1855:
1825:
1729:Falconer's formula
1721:assortative mating
1716:
1687:Sibling comparison
1664:
1617:
1375:
1259:
1238:
1202:
1166:
1106:
990:
969:
933:
897:
833:
831:
530:
489:for narrow sense.
468:
353:
272:
106:selective breeding
104:, particularly in
47:
6923:
6922:
6905:Pavlovian session
6777:Nikita Khrushchev
6719:
6718:
6652:Eric F. Wieschaus
6614:
6613:
6432:Pattern formation
6336:Fitness landscape
6198:
6197:
6148:Genetic genealogy
6143:Fitness landscape
5872:
5871:
5694:978-0-87893-481-2
5650:978-1-317-60590-4
5634:Joseph J (2014).
5340:978-0-8147-8723-6
5298:978-1-898059-47-9
5284:The Gene Illusion
5277:Joseph J (2004).
5219:978-0-582-24302-6
5192:978-0-7167-2056-0
4955:Behavior Genetics
4877:Behavior Genetics
4593:978-0-87893-308-2
3988:978-0-393-26313-8
3886:molecular-genetic
3834:
3833:
3585:genetic dominance
3573:reverse causality
3506:
3505:
3491:
3463:
3435:{\displaystyle 0}
3409:
3376:
3348:
3320:{\displaystyle 0}
3294:
3266:{\displaystyle 0}
3240:
3223:Parent-Offspring
3212:{\displaystyle 1}
3191:{\displaystyle 1}
3144:{\displaystyle r}
3040:{\displaystyle r}
2900:{\displaystyle e}
2698:statistical power
2677:
2624:
2541:
2524:
2523:
2486:
2463:{\displaystyle W}
2382:
2339:
2316:{\displaystyle S}
2190:{\displaystyle r}
2170:{\displaystyle n}
2123:
2093:
2065:
1970:within sire group
1932:
1885:{\displaystyle e}
1734:=2(r(MZ)-r(DZ)).
1615:
1602:
1486:was developed by
1458:was developed by
1456:school of thought
1389:linear regression
773:
711:
703:
466:
351:
208:) + Environment (
126:genetic variation
110:behavior genetics
78:genetic variation
16:(Redirected from
6953:
6746:
6739:
6732:
6723:
6662:William McGinnis
6631:Richard Lewontin
6626:C. H. Waddington
6498:
6475:Neutral networks
6225:
6218:
6211:
6202:
6107:J. B. S. Haldane
5899:
5892:
5885:
5876:
5788:
5781:
5774:
5765:
5735:
5698:
5669:
5668:
5666:
5665:
5659:
5642:
5631:
5625:
5624:
5613:10.1002/hast.496
5596:
5590:
5589:
5572:(3–4): 227–241.
5561:
5552:
5549:
5543:
5542:
5540:
5538:
5482:
5476:
5475:
5462:
5430:
5424:
5423:
5412:10.1002/wcs.1400
5395:
5389:
5388:
5386:
5384:
5362:
5356:
5355:
5353:
5352:
5320:
5314:
5313:
5311:
5310:
5274:
5268:
5267:
5257:
5230:
5224:
5223:
5203:
5197:
5196:
5176:
5170:
5169:
5168:
5162:
5152:
5142:
5118:
5112:
5111:
5101:
5091:
5059:
5053:
5052:
5050:
5049:
5043:
5004:
4995:
4989:
4988:
4978:
4946:
4940:
4939:
4924:(4th ed.).
4915:
4909:
4908:
4872:
4866:
4865:
4854:10.1037/h0043487
4837:
4831:
4830:
4820:
4788:
4782:
4781:
4771:
4761:
4729:
4723:
4722:
4720:
4719:
4699:
4693:
4692:
4676:
4665:
4664:
4662:
4660:
4654:
4615:
4606:
4600:
4597:
4574:
4572:
4549:Wills C (2007).
4546:
4540:
4539:
4503:
4497:
4496:
4486:
4454:
4448:
4447:
4445:
4444:
4438:
4399:
4386:
4375:
4374:
4338:
4332:
4331:
4329:
4328:
4322:
4291:
4282:
4276:
4275:
4273:
4272:
4246:
4214:
4208:
4207:
4205:
4204:
4198:
4175:
4149:
4140:
4131:
4130:
4128:
4127:
4121:
4091:(9): 1304–1310.
4082:
4073:
4064:
4063:
4053:
4021:
4015:
4014:
4008:
4000:
3974:
3968:
3967:
3965:
3963:
3948:Nature Education
3939:
3829:
3826:
3820:
3796:
3795:
3788:
3775:
3773:
3772:
3767:
3756:
3745:
3731:
3699:
3697:
3696:
3691:
3686:
3685:
3656:
3654:
3653:
3648:
3643:
3642:
3577:maternal effects
3531:Bayesian methods
3502:
3500:
3499:
3494:
3492:
3484:
3474:
3472:
3471:
3466:
3464:
3456:
3441:
3439:
3438:
3433:
3420:
3418:
3417:
3412:
3410:
3402:
3387:
3385:
3384:
3379:
3377:
3369:
3359:
3357:
3356:
3351:
3349:
3341:
3326:
3324:
3323:
3318:
3305:
3303:
3302:
3297:
3295:
3287:
3272:
3270:
3269:
3264:
3251:
3249:
3248:
3243:
3241:
3233:
3218:
3216:
3215:
3210:
3197:
3195:
3194:
3189:
3176:Identical Twins
3171:
3169:
3168:
3163:
3150:
3148:
3147:
3142:
3126:
3116:
3114:
3113:
3108:
3091:
3089:
3088:
3083:
3066:
3064:
3063:
3058:
3046:
3044:
3043:
3038:
3023:
3021:
3020:
3015:
3010:
3009:
2994:
2993:
2966:linear functions
2963:
2961:
2960:
2955:
2953:
2952:
2936:
2934:
2933:
2928:
2926:
2925:
2906:
2904:
2903:
2898:
2886:
2884:
2883:
2878:
2876:
2875:
2856:
2854:
2853:
2848:
2846:
2845:
2829:
2827:
2826:
2821:
2819:
2818:
2797:
2795:
2794:
2789:
2778:
2777:
2762:
2761:
2749:
2748:
2730:
2729:
2688:
2686:
2685:
2680:
2678:
2676:
2650:
2630:
2625:
2623:
2622:
2621:
2609:
2608:
2598:
2597:
2588:
2583:
2582:
2562:
2560:
2559:
2554:
2552:
2551:
2542:
2534:
2520:
2518:
2517:
2512:
2510:
2509:
2497:
2496:
2487:
2479:
2469:
2467:
2466:
2461:
2448:
2446:
2445:
2440:
2407:
2405:
2404:
2399:
2394:
2393:
2392:
2383:
2375:
2363:
2362:
2350:
2349:
2340:
2332:
2322:
2320:
2319:
2314:
2301:
2299:
2298:
2293:
2257:
2250:
2248:
2247:
2242:
2240:
2239:
2223:
2221:
2220:
2215:
2213:
2212:
2196:
2194:
2193:
2188:
2176:
2174:
2173:
2168:
2144:
2142:
2141:
2136:
2134:
2133:
2124:
2116:
2108:
2094:
2086:
2066:
2058:
2047:
2027:
2010:
1981:among sire group
1959:
1957:
1956:
1951:
1943:
1942:
1933:
1925:
1914:
1913:
1891:
1889:
1888:
1883:
1864:
1862:
1861:
1856:
1854:
1853:
1834:
1832:
1831:
1826:
1818:
1817:
1799:
1798:
1626:
1624:
1623:
1618:
1616:
1608:
1603:
1595:
1590:
1589:
1497:and expanded at
1384:
1382:
1381:
1376:
1368:
1367:
1337:
1336:
1306:
1305:
1268:
1266:
1265:
1260:
1254:
1249:
1218:
1213:
1182:
1177:
1138:
1115:
1113:
1112:
1107:
1099:
1098:
1068:
1067:
1037:
1036:
999:
997:
996:
991:
985:
980:
949:
944:
913:
908:
869:
847:weighted average
842:
840:
839:
834:
832:
819:
818:
809:
808:
790:
789:
774:
771:
760:
759:
747:
746:
728:
727:
712:
709:
704:
701:
693:
686:
685:
676:
675:
653:
652:
640:
639:
610:
609:
477:
475:
474:
469:
467:
465:
455:
443:
433:
421:
416:
415:
362:
360:
359:
354:
352:
350:
340:
328:
318:
306:
301:
300:
281:
279:
278:
273:
268:
267:
70:phenotypic trait
21:
6961:
6960:
6956:
6955:
6954:
6952:
6951:
6950:
6926:
6925:
6924:
6919:
6888:Soviet policies
6883:
6837:
6833:Nikolai Vavilov
6813:Zhores Medvedev
6803:Wacław Gajewski
6791:
6755:
6750:
6720:
6715:
6706:
6685:
6672:Sean B. Carroll
6610:
6542:
6489:
6453:
6405:
6386:Maternal effect
6369:
6302:
6239:
6229:
6199:
6194:
6181:
6116:
6090:
6052:
6036:
6034:
6027:
5994:
5925:Genetic linkage
5908:
5903:
5873:
5868:
5837:
5801:
5792:
5759:Wayback Machine
5743:
5738:
5716:10.1002/per.835
5701:
5695:
5682:
5678:
5676:Further reading
5673:
5672:
5663:
5661:
5657:
5651:
5640:
5633:
5632:
5628:
5598:
5597:
5593:
5563:
5562:
5555:
5550:
5546:
5536:
5534:
5484:
5483:
5479:
5432:
5431:
5427:
5397:
5396:
5392:
5382:
5380:
5364:
5363:
5359:
5350:
5348:
5341:
5322:
5321:
5317:
5308:
5306:
5299:
5291:. p. 141.
5276:
5275:
5271:
5232:
5231:
5227:
5220:
5205:
5204:
5200:
5193:
5178:
5177:
5173:
5163:
5133:(2): e1000008.
5120:
5119:
5115:
5074:(27): 7377–82.
5061:
5060:
5056:
5047:
5045:
5041:
5002:
4997:
4996:
4992:
4948:
4947:
4943:
4936:
4917:
4916:
4912:
4874:
4873:
4869:
4839:
4838:
4834:
4790:
4789:
4785:
4731:
4730:
4726:
4717:
4715:
4701:
4700:
4696:
4678:
4677:
4668:
4658:
4656:
4652:
4613:
4608:
4607:
4603:
4594:
4581:
4559:(Book Review).
4548:
4547:
4543:
4505:
4504:
4500:
4456:
4455:
4451:
4442:
4440:
4436:
4414:10.1038/nrg2322
4397:
4388:
4387:
4378:
4340:
4339:
4335:
4326:
4324:
4320:
4289:
4284:
4283:
4279:
4270:
4268:
4216:
4215:
4211:
4202:
4200:
4196:
4152:Nature Genetics
4147:
4142:
4141:
4134:
4125:
4123:
4119:
4097:10.1038/ng.3941
4085:Nature Genetics
4080:
4075:
4074:
4067:
4023:
4022:
4018:
4001:
3989:
3976:
3975:
3971:
3961:
3959:
3941:
3940:
3936:
3931:
3909:
3854:social sciences
3846:Richard Bentall
3830:
3824:
3821:
3810:
3797:
3793:
3786:
3716:
3715:
3677:
3666:
3665:
3634:
3626:
3625:
3606:
3593:
3511:
3478:
3477:
3450:
3449:
3424:
3423:
3396:
3395:
3363:
3362:
3335:
3334:
3309:
3308:
3281:
3280:
3255:
3254:
3227:
3226:
3201:
3200:
3180:
3179:
3154:
3153:
3133:
3132:
3096:
3095:
3071:
3070:
3049:
3048:
3029:
3028:
3001:
2985:
2974:
2973:
2944:
2939:
2938:
2917:
2912:
2911:
2889:
2888:
2864:
2859:
2858:
2837:
2832:
2831:
2810:
2805:
2804:
2766:
2753:
2740:
2718:
2713:
2712:
2706:
2651:
2631:
2613:
2600:
2599:
2589:
2574:
2569:
2568:
2543:
2528:
2527:
2501:
2488:
2473:
2472:
2452:
2451:
2416:
2415:
2384:
2354:
2341:
2326:
2325:
2305:
2304:
2278:
2277:
2231:
2226:
2225:
2204:
2199:
2198:
2179:
2178:
2159:
2158:
2155:
2125:
2101:
2020:
1992:
1991:
1966:
1934:
1905:
1900:
1899:
1874:
1873:
1870:
1845:
1840:
1839:
1809:
1790:
1785:
1784:
1778:
1771:
1763:
1708:
1702:
1689:
1652:
1581:
1576:
1575:
1569:
1561:
1533:identical twins
1414:
1397:
1356:
1325:
1294:
1274:
1273:
1123:
1122:
1087:
1056:
1025:
1005:
1004:
854:
853:
830:
829:
810:
800:
778:
751:
738:
716:
702:Population mean
691:
690:
677:
667:
644:
631:
611:
598:
589:
588:
584:
577:
566:
559:
552:
545:
522:
444:
422:
407:
402:
401:
329:
307:
292:
287:
286:
259:
254:
253:
190:
150:transcriptional
122:
112:(for instance,
76:that is due to
35:
28:
23:
22:
18:Heritable trait
15:
12:
11:
5:
6959:
6957:
6949:
6948:
6943:
6938:
6928:
6927:
6921:
6920:
6918:
6917:
6912:
6907:
6902:
6897:
6891:
6889:
6885:
6884:
6882:
6881:
6876:
6871:
6866:
6861:
6856:
6851:
6845:
6843:
6839:
6838:
6836:
6835:
6830:
6825:
6820:
6818:Georgii Nadson
6815:
6810:
6805:
6799:
6797:
6793:
6792:
6790:
6789:
6784:
6779:
6774:
6769:
6767:Trofim Lysenko
6763:
6761:
6757:
6756:
6751:
6749:
6748:
6741:
6734:
6726:
6717:
6716:
6711:
6708:
6707:
6705:
6704:
6699:
6693:
6691:
6687:
6686:
6684:
6683:
6682:
6681:
6669:
6664:
6659:
6654:
6649:
6648:
6647:
6636:François Jacob
6633:
6628:
6622:
6620:
6616:
6615:
6612:
6611:
6609:
6608:
6603:
6598:
6593:
6588:
6583:
6578:
6573:
6572:
6571:
6561:
6556:
6550:
6548:
6544:
6543:
6541:
6540:
6535:
6530:
6525:
6520:
6515:
6510:
6504:
6502:
6495:
6491:
6490:
6488:
6487:
6482:
6477:
6472:
6467:
6461:
6459:
6455:
6454:
6452:
6451:
6446:
6441:
6436:
6435:
6434:
6429:
6419:
6413:
6411:
6407:
6406:
6404:
6403:
6398:
6393:
6388:
6383:
6377:
6375:
6371:
6370:
6368:
6367:
6365:Sequence space
6362:
6357:
6352:
6347:
6342:
6333:
6328:
6323:
6318:
6312:
6310:
6304:
6303:
6301:
6300:
6295:
6294:
6293:
6283:
6278:
6273:
6268:
6263:
6258:
6253:
6247:
6245:
6241:
6240:
6230:
6228:
6227:
6220:
6213:
6205:
6196:
6195:
6193:
6192:
6186:
6183:
6182:
6180:
6179:
6174:
6172:Phylogeography
6169:
6164:
6162:Microevolution
6159:
6150:
6145:
6140:
6135:
6130:
6124:
6122:
6121:Related topics
6118:
6117:
6115:
6114:
6109:
6104:
6098:
6096:
6092:
6091:
6089:
6088:
6083:
6078:
6076:Founder effect
6073:
6068:
6062:
6060:
6054:
6053:
6051:
6050:
6045:
6039:
6037:
6032:
6029:
6028:
6026:
6025:
6020:
6015:
6010:
6004:
6002:
5996:
5995:
5993:
5992:
5987:
5982:
5977:
5972:
5967:
5962:
5957:
5955:Price equation
5952:
5947:
5945:Neutral theory
5942:
5937:
5932:
5927:
5922:
5916:
5914:
5910:
5909:
5904:
5902:
5901:
5894:
5887:
5879:
5870:
5869:
5867:
5866:
5861:
5856:
5851:
5845:
5843:
5842:Related Topics
5839:
5838:
5836:
5835:
5830:
5828:Candidate gene
5825:
5820:
5815:
5809:
5807:
5803:
5802:
5793:
5791:
5790:
5783:
5776:
5768:
5762:
5761:
5749:
5742:
5741:External links
5739:
5737:
5736:
5710:(4): 287–294.
5699:
5693:
5679:
5677:
5674:
5671:
5670:
5649:
5626:
5591:
5553:
5544:
5477:
5425:
5406:(1–2): e1400.
5390:
5357:
5339:
5315:
5297:
5269:
5225:
5218:
5198:
5191:
5171:
5113:
5054:
5013:(1): 109–120.
4990:
4941:
4935:978-0582243026
4934:
4910:
4867:
4832:
4783:
4724:
4704:"Heritability"
4694:
4666:
4624:(5): 160–164.
4601:
4599:
4598:
4592:
4541:
4498:
4449:
4376:
4333:
4300:(3): 244–260.
4277:
4229:(2–3): 89–96.
4209:
4158:(8): 859–864.
4132:
4065:
4016:
3987:
3969:
3933:
3932:
3930:
3927:
3926:
3925:
3920:
3915:
3908:
3905:
3832:
3831:
3800:
3798:
3791:
3785:
3782:
3765:
3762:
3759:
3755:
3752:
3749:
3744:
3740:
3737:
3734:
3730:
3727:
3724:
3701:
3700:
3689:
3684:
3680:
3676:
3673:
3646:
3641:
3637:
3633:
3605:
3602:
3592:
3589:
3510:
3507:
3504:
3503:
3490:
3487:
3475:
3462:
3459:
3447:
3443:
3442:
3431:
3421:
3408:
3405:
3393:
3392:First Cousins
3389:
3388:
3375:
3372:
3360:
3347:
3344:
3332:
3331:Full Siblings
3328:
3327:
3316:
3306:
3293:
3290:
3278:
3277:Half Siblings
3274:
3273:
3262:
3252:
3239:
3236:
3224:
3220:
3219:
3208:
3198:
3187:
3177:
3173:
3172:
3161:
3151:
3140:
3130:
3106:
3103:
3081:
3078:
3056:
3036:
3025:
3024:
3013:
3008:
3004:
3000:
2997:
2992:
2988:
2984:
2981:
2951:
2947:
2924:
2920:
2896:
2874:
2871:
2867:
2844:
2840:
2817:
2813:
2799:
2798:
2787:
2784:
2781:
2776:
2773:
2769:
2765:
2760:
2756:
2752:
2747:
2743:
2739:
2736:
2733:
2728:
2725:
2721:
2705:
2702:
2675:
2672:
2669:
2666:
2663:
2660:
2657:
2654:
2649:
2646:
2643:
2640:
2637:
2634:
2628:
2620:
2616:
2612:
2607:
2603:
2596:
2592:
2586:
2581:
2577:
2550:
2546:
2540:
2537:
2522:
2521:
2508:
2504:
2500:
2495:
2491:
2485:
2482:
2470:
2459:
2449:
2438:
2435:
2432:
2429:
2426:
2423:
2413:
2409:
2408:
2397:
2391:
2387:
2381:
2378:
2372:
2369:
2366:
2361:
2357:
2353:
2348:
2344:
2338:
2335:
2323:
2312:
2302:
2291:
2288:
2285:
2275:
2271:
2270:
2267:
2264:
2261:
2238:
2234:
2211:
2207:
2186:
2166:
2154:
2151:
2147:
2146:
2132:
2128:
2122:
2119:
2114:
2111:
2107:
2104:
2100:
2097:
2092:
2089:
2084:
2081:
2078:
2075:
2072:
2069:
2064:
2061:
2056:
2053:
2050:
2046:
2043:
2040:
2037:
2033:
2030:
2026:
2023:
2019:
2016:
2013:
2009:
2006:
2003:
2000:
1965:
1962:
1961:
1960:
1949:
1946:
1941:
1937:
1931:
1928:
1923:
1920:
1917:
1912:
1908:
1881:
1868:
1852:
1848:
1836:
1835:
1824:
1821:
1816:
1812:
1808:
1805:
1802:
1797:
1793:
1776:
1770:
1767:
1762:
1759:
1704:Main article:
1701:
1698:
1688:
1685:
1660:Francis Galton
1651:
1648:
1628:
1627:
1614:
1611:
1606:
1601:
1598:
1593:
1588:
1584:
1568:
1565:
1560:
1557:
1438:standard error
1413:
1410:
1396:
1393:
1374:
1371:
1366:
1363:
1359:
1355:
1352:
1349:
1346:
1343:
1340:
1335:
1332:
1328:
1324:
1321:
1318:
1315:
1312:
1309:
1304:
1301:
1297:
1293:
1290:
1287:
1284:
1281:
1270:
1269:
1258:
1253:
1248:
1245:
1241:
1237:
1234:
1231:
1228:
1225:
1222:
1217:
1212:
1209:
1205:
1201:
1198:
1195:
1192:
1189:
1186:
1181:
1176:
1173:
1169:
1165:
1162:
1159:
1156:
1153:
1150:
1147:
1144:
1141:
1137:
1134:
1131:
1105:
1102:
1097:
1094:
1090:
1086:
1083:
1080:
1077:
1074:
1071:
1066:
1063:
1059:
1055:
1052:
1049:
1046:
1043:
1040:
1035:
1032:
1028:
1024:
1021:
1018:
1015:
1012:
1001:
1000:
989:
984:
979:
976:
972:
968:
965:
962:
959:
956:
953:
948:
943:
940:
936:
932:
929:
926:
923:
920:
917:
912:
907:
904:
900:
896:
893:
890:
887:
884:
881:
878:
875:
872:
868:
865:
862:
828:
825:
822:
817:
813:
807:
803:
799:
796:
793:
788:
785:
781:
777:
769:
766:
763:
758:
754:
750:
745:
741:
737:
734:
731:
726:
723:
719:
715:
707:
699:
696:
694:
692:
689:
684:
680:
674:
670:
666:
662:
659:
656:
651:
647:
643:
638:
634:
630:
626:
623:
620:
617:
614:
612:
608:
605:
601:
597:
596:
582:
575:
564:
557:
550:
543:
535:The number of
521:
518:
481:An upper case
479:
478:
464:
461:
458:
454:
451:
448:
442:
439:
436:
432:
429:
426:
419:
414:
410:
364:
363:
349:
346:
343:
339:
336:
333:
327:
324:
321:
317:
314:
311:
304:
299:
295:
282:is defined as
271:
266:
262:
242:
241:
214:
213:
204:) = Genotype (
189:
186:
121:
118:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
6958:
6947:
6944:
6942:
6939:
6937:
6934:
6933:
6931:
6916:
6913:
6911:
6908:
6906:
6903:
6901:
6898:
6896:
6893:
6892:
6890:
6886:
6880:
6879:Vernalization
6877:
6875:
6872:
6870:
6867:
6865:
6862:
6860:
6859:Hybridization
6857:
6855:
6852:
6850:
6847:
6846:
6844:
6840:
6834:
6831:
6829:
6826:
6824:
6821:
6819:
6816:
6814:
6811:
6809:
6806:
6804:
6801:
6800:
6798:
6794:
6788:
6785:
6783:
6782:Joseph Stalin
6780:
6778:
6775:
6773:
6770:
6768:
6765:
6764:
6762:
6758:
6754:
6747:
6742:
6740:
6735:
6733:
6728:
6727:
6724:
6714:
6709:
6703:
6700:
6698:
6695:
6694:
6692:
6688:
6680:
6679:
6675:
6674:
6673:
6670:
6668:
6665:
6663:
6660:
6658:
6655:
6653:
6650:
6646:
6643:
6642:
6641:
6640:Jacques Monod
6637:
6634:
6632:
6629:
6627:
6624:
6623:
6621:
6617:
6607:
6604:
6602:
6599:
6597:
6594:
6592:
6589:
6587:
6584:
6582:
6579:
6577:
6574:
6570:
6567:
6566:
6565:
6562:
6560:
6557:
6555:
6554:Homeotic gene
6552:
6551:
6549:
6545:
6539:
6536:
6534:
6531:
6529:
6526:
6524:
6521:
6519:
6516:
6514:
6511:
6509:
6506:
6505:
6503:
6499:
6496:
6492:
6486:
6483:
6481:
6478:
6476:
6473:
6471:
6468:
6466:
6463:
6462:
6460:
6456:
6450:
6447:
6445:
6442:
6440:
6437:
6433:
6430:
6428:
6425:
6424:
6423:
6422:Morphogenesis
6420:
6418:
6415:
6414:
6412:
6408:
6402:
6399:
6397:
6394:
6392:
6389:
6387:
6384:
6382:
6379:
6378:
6376:
6372:
6366:
6363:
6361:
6358:
6356:
6353:
6351:
6348:
6346:
6343:
6341:
6337:
6334:
6332:
6329:
6327:
6324:
6322:
6319:
6317:
6314:
6313:
6311:
6309:
6305:
6299:
6296:
6292:
6289:
6288:
6287:
6284:
6282:
6279:
6277:
6274:
6272:
6269:
6267:
6264:
6262:
6259:
6257:
6256:Reaction norm
6254:
6252:
6249:
6248:
6246:
6242:
6238:
6234:
6226:
6221:
6219:
6214:
6212:
6207:
6206:
6203:
6191:
6188:
6187:
6184:
6178:
6175:
6173:
6170:
6168:
6165:
6163:
6160:
6158:
6154:
6151:
6149:
6146:
6144:
6141:
6139:
6136:
6134:
6131:
6129:
6126:
6125:
6123:
6119:
6113:
6112:Sewall Wright
6110:
6108:
6105:
6103:
6100:
6099:
6097:
6093:
6087:
6084:
6082:
6079:
6077:
6074:
6072:
6069:
6067:
6064:
6063:
6061:
6059:
6058:Genetic drift
6055:
6049:
6046:
6044:
6041:
6040:
6038:
6030:
6024:
6021:
6019:
6016:
6014:
6011:
6009:
6006:
6005:
6003:
6001:
5997:
5991:
5988:
5986:
5983:
5981:
5978:
5976:
5973:
5971:
5968:
5966:
5963:
5961:
5958:
5956:
5953:
5951:
5948:
5946:
5943:
5941:
5938:
5936:
5933:
5931:
5928:
5926:
5923:
5921:
5918:
5917:
5915:
5911:
5907:
5900:
5895:
5893:
5888:
5886:
5881:
5880:
5877:
5865:
5862:
5860:
5857:
5855:
5852:
5850:
5847:
5846:
5844:
5840:
5834:
5831:
5829:
5826:
5824:
5821:
5819:
5816:
5814:
5811:
5810:
5808:
5804:
5800:
5796:
5789:
5784:
5782:
5777:
5775:
5770:
5769:
5766:
5760:
5756:
5753:
5750:
5748:
5745:
5744:
5740:
5733:
5729:
5725:
5721:
5717:
5713:
5709:
5705:
5700:
5696:
5690:
5686:
5681:
5680:
5675:
5656:
5652:
5646:
5639:
5638:
5630:
5627:
5622:
5618:
5614:
5610:
5606:
5602:
5595:
5592:
5587:
5583:
5579:
5575:
5571:
5567:
5560:
5558:
5554:
5548:
5545:
5532:
5528:
5524:
5520:
5516:
5512:
5508:
5504:
5500:
5496:
5492:
5488:
5481:
5478:
5474:
5470:
5466:
5461:
5456:
5452:
5448:
5444:
5440:
5436:
5429:
5426:
5421:
5417:
5413:
5409:
5405:
5401:
5394:
5391:
5378:
5374:
5373:
5372:The Telegraph
5368:
5361:
5358:
5346:
5342:
5336:
5332:
5328:
5327:
5319:
5316:
5304:
5300:
5294:
5290:
5286:
5285:
5280:
5273:
5270:
5265:
5261:
5256:
5251:
5247:
5243:
5239:
5236:(June 2006).
5235:
5229:
5226:
5221:
5215:
5211:
5210:
5202:
5199:
5194:
5188:
5184:
5183:
5175:
5172:
5167:
5160:
5156:
5151:
5146:
5141:
5136:
5132:
5128:
5127:PLOS Genetics
5124:
5117:
5114:
5109:
5105:
5100:
5095:
5090:
5085:
5081:
5077:
5073:
5069:
5065:
5058:
5055:
5040:
5036:
5032:
5028:
5024:
5020:
5016:
5012:
5008:
5001:
4994:
4991:
4986:
4982:
4977:
4972:
4968:
4964:
4960:
4956:
4952:
4945:
4942:
4937:
4931:
4927:
4923:
4922:
4914:
4911:
4906:
4902:
4898:
4894:
4890:
4886:
4883:(5): 467–73.
4882:
4878:
4871:
4868:
4863:
4859:
4855:
4851:
4848:(6): 353–72.
4847:
4843:
4836:
4833:
4828:
4824:
4819:
4814:
4810:
4806:
4802:
4798:
4794:
4787:
4784:
4779:
4775:
4770:
4765:
4760:
4755:
4751:
4747:
4744:(4): 1193–8.
4743:
4739:
4735:
4728:
4725:
4713:
4709:
4705:
4698:
4695:
4690:
4686:
4682:
4675:
4673:
4671:
4667:
4651:
4647:
4643:
4639:
4635:
4631:
4627:
4623:
4619:
4612:
4605:
4602:
4595:
4589:
4585:
4579:
4576:
4575:
4571:
4566:
4562:
4558:
4557:
4552:
4545:
4542:
4537:
4533:
4529:
4525:
4521:
4517:
4514:(2): 99–128.
4513:
4509:
4502:
4499:
4494:
4490:
4485:
4480:
4476:
4472:
4468:
4464:
4460:
4453:
4450:
4435:
4431:
4427:
4423:
4419:
4415:
4411:
4408:(4): 255–66.
4407:
4403:
4396:
4392:
4389:Visscher PM,
4385:
4383:
4381:
4377:
4372:
4368:
4364:
4360:
4356:
4352:
4348:
4344:
4337:
4334:
4319:
4315:
4311:
4307:
4303:
4299:
4295:
4288:
4281:
4278:
4266:
4262:
4258:
4254:
4250:
4245:
4244:2027.42/42804
4240:
4236:
4232:
4228:
4224:
4220:
4213:
4210:
4195:
4191:
4187:
4183:
4179:
4174:
4169:
4165:
4161:
4157:
4153:
4146:
4139:
4137:
4133:
4118:
4114:
4110:
4106:
4102:
4098:
4094:
4090:
4086:
4079:
4072:
4070:
4066:
4061:
4057:
4052:
4047:
4043:
4039:
4035:
4031:
4027:
4020:
4017:
4012:
4006:
3998:
3994:
3990:
3984:
3980:
3973:
3970:
3957:
3953:
3949:
3945:
3938:
3935:
3928:
3924:
3921:
3919:
3916:
3914:
3911:
3910:
3906:
3904:
3902:
3898:
3896:
3891:
3887:
3883:
3878:
3876:
3872:
3867:
3863:
3859:
3855:
3851:
3847:
3843:
3839:
3828:
3818:
3814:
3808:
3806:
3801:This section
3799:
3790:
3789:
3784:Controversies
3783:
3781:
3779:
3760:
3742:
3735:
3712:
3709:
3705:
3687:
3682:
3678:
3674:
3671:
3664:
3663:
3662:
3660:
3639:
3635:
3623:
3615:
3610:
3603:
3601:
3599:
3590:
3588:
3586:
3582:
3578:
3574:
3569:
3567:
3564:
3560:
3557:
3554:
3551:
3547:
3543:
3539:
3536:
3532:
3528:
3523:
3521:
3517:
3508:
3488:
3485:
3476:
3460:
3457:
3448:
3445:
3444:
3429:
3422:
3406:
3403:
3394:
3391:
3390:
3373:
3370:
3361:
3345:
3342:
3333:
3330:
3329:
3314:
3307:
3291:
3288:
3279:
3276:
3275:
3260:
3253:
3237:
3234:
3225:
3222:
3221:
3206:
3199:
3185:
3178:
3175:
3174:
3159:
3152:
3138:
3131:
3129:Relationship
3128:
3127:
3121:
3118:
3104:
3101:
3093:
3079:
3076:
3068:
3067:are found as
3054:
3034:
3011:
3006:
3002:
2998:
2995:
2990:
2986:
2982:
2979:
2971:
2970:
2969:
2967:
2949:
2945:
2922:
2918:
2908:
2894:
2872:
2869:
2865:
2842:
2838:
2815:
2811:
2802:
2785:
2782:
2779:
2774:
2771:
2767:
2763:
2758:
2754:
2750:
2745:
2741:
2737:
2734:
2731:
2726:
2723:
2719:
2711:
2710:
2709:
2703:
2701:
2699:
2695:
2690:
2673:
2667:
2664:
2661:
2655:
2652:
2644:
2641:
2638:
2632:
2626:
2618:
2614:
2610:
2605:
2601:
2594:
2590:
2584:
2579:
2575:
2566:
2548:
2544:
2538:
2535:
2506:
2502:
2498:
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6569:eyeless gene
6465:Evolvability
6439:Segmentation
6316:Canalisation
6286:Heterochrony
6276:Heritability
6275:
6244:Key concepts
6128:Biogeography
6102:R. A. Fisher
5980:Heritability
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5913:Key concepts
5813:Heritability
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3882:twin studies
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114:twin studies
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6828:Tan Jiazhen
6823:Hans Stubbe
6760:Lysenkoists
6667:Mike Levine
6576:Distal-less
6401:Polyphenism
6381:Epigenetics
6233:development
6081:Coalescence
5279:"Chapter 5"
4349:(1): 1–27.
4173:10616/47311
3866:David Shenk
3862:David Moore
3838:Steven Rose
3825:August 2016
1769:Basic model
1725:correlation
1694:familiarity
1422:statistical
1416:Since only
1395:Assumptions
569:interaction
200:Phenotype (
43:differences
6930:Categories
6869:Lamarckism
6796:Dissidents
6753:Lysenkoism
6645:Lac operon
6470:Robustness
6449:Modularity
6444:Metamerism
6350:Plasticity
6345:Pleiotropy
6298:Heterotopy
6023:Ecological
6013:Artificial
5664:2016-04-02
5351:2016-04-02
5309:2016-04-02
5048:2020-09-06
4803:: 422–39.
4718:2020-02-20
4659:29 October
4563:(4): 382.
4443:2015-08-28
4327:2019-12-05
4271:2020-12-24
4203:2021-02-08
4126:2020-09-06
3929:References
3842:Jay Joseph
3805:unbalanced
3529:(REML) or
2177:sires and
1985:error term
1757:=1-r(MZ).
1706:Twin study
1669:regression
1658:Figure 2.
1538:twin study
1476:J. L. Lush
232:) + 2 Cov(
188:Definition
178:antibiotic
166:hair color
162:univariate
135:inbreeding
74:population
6596:Morphogen
6581:Engrailed
6564:Pax genes
6485:Tinkering
6331:Epistasis
6326:Dominance
6237:phenotype
6133:Evolution
6000:Selection
5818:Dominance
5724:0890-2070
5035:143217984
5027:1469-1825
4689:422371269
4638:0963-7214
4578:review of
4536:204981536
4508:Cognition
4314:1878-7304
4190:220260262
4005:cite book
3997:908409996
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3817:talk page
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170:eye color
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6842:Concepts
6787:VASKhNIL
6559:Hox gene
6547:Elements
6528:Homeobox
6157:genomics
6095:Founders
5864:Heredity
5854:Genomics
5795:Genetics
5755:Archived
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5655:Archived
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1468:C. C. Li
571:between
371:dominant
228:) + Var(
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142:canalize
120:Overview
62:genetics
58:breeding
32:heredity
6690:Debates
6501:Systems
6427:Eyespot
6291:Neoteny
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5975:Fitness
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5499:Bibcode
5491:Science
5460:5812976
5383:4 April
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2263:d.f.
1974:error
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1434:twins
394:locus
130:genes
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5720:ISSN
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5617:PMID
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5385:2018
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4685:OCLC
4661:2013
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4310:ISSN
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3993:OCLC
3983:ISBN
3964:2015
3873:and
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1872:and
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3374:4
3371:1
3346:2
3343:1
3315:0
3292:4
3289:1
3261:0
3238:2
3235:1
3207:1
3186:1
3139:r
3105:=
3080:=
3077:r
3035:r
3012:,
3007:d
3003:V
2996:+
2991:a
2987:V
2983:r
2980:=
2950:d
2946:V
2923:a
2919:V
2895:e
2873:j
2870:i
2866:d
2843:j
2816:i
2786:,
2783:e
2780:+
2775:j
2772:i
2768:d
2764:+
2759:j
2751:+
2746:i
2738:+
2732:=
2727:j
2724:i
2720:y
2674:W
2671:)
2668:1
2662:r
2659:(
2656:+
2653:S
2648:)
2645:W
2639:S
2636:(
2633:4
2627:=
2619:e
2615:V
2611:+
2606:g
2602:V
2595:g
2591:V
2585:=
2580:2
2576:H
2549:g
2545:V
2539:4
2536:1
2507:e
2503:V
2499:+
2494:g
2490:V
2484:4
2481:3
2458:W
2437:)
2434:1
2428:r
2425:(
2422:n
2396:)
2390:g
2386:V
2380:4
2377:1
2371:(
2368:r
2365:+
2360:e
2356:V
2352:+
2347:g
2343:V
2337:4
2334:3
2311:S
2290:1
2284:n
2237:e
2233:V
2210:g
2206:V
2185:r
2165:n
2145:,
2131:g
2127:V
2121:4
2118:1
2113:=
2110:)
2103:e
2099:+
2096:g
2091:2
2088:1
2083:+
2077:,
2074:e
2071:+
2068:g
2063:2
2060:1
2055:+
2049:(
2045:r
2042:r
2039:o
2036:c
2032:=
2029:)
2022:z
2018:,
2015:z
2012:(
2008:r
2005:r
2002:o
1999:c
1948:e
1945:+
1940:i
1936:g
1930:2
1927:1
1922:+
1916:=
1911:i
1907:z
1880:e
1869:i
1867:G
1851:i
1847:g
1823:e
1820:+
1815:i
1811:g
1807:+
1801:=
1796:i
1792:y
1777:i
1775:G
1755:e
1751:e
1747:h
1743:c
1739:c
1732:H
1644:t
1640:b
1632:r
1613:r
1610:t
1605:=
1600:r
1597:b
1592:=
1587:2
1583:h
1478:(
1470:(
1418:P
1400:"
1370:=
1365:B
1362:B
1358:d
1354:)
1351:B
1348:B
1345:(
1342:f
1339:+
1334:b
1331:B
1327:d
1323:)
1320:b
1317:B
1314:(
1311:f
1308:+
1303:b
1300:b
1296:d
1292:)
1289:b
1286:b
1283:(
1280:f
1257:,
1252:2
1247:B
1244:B
1240:d
1236:)
1233:B
1230:B
1227:(
1224:f
1221:+
1216:2
1211:b
1208:B
1204:d
1200:)
1197:b
1194:B
1191:(
1188:f
1185:+
1180:2
1175:b
1172:b
1168:d
1164:)
1161:b
1158:b
1155:(
1152:f
1149:=
1146:)
1143:D
1140:(
1136:r
1133:a
1130:V
1101:=
1096:B
1093:B
1089:a
1085:)
1082:B
1079:B
1076:(
1073:f
1070:+
1065:b
1062:B
1058:a
1054:)
1051:b
1048:B
1045:(
1042:f
1039:+
1034:b
1031:b
1027:a
1023:)
1020:b
1017:b
1014:(
1011:f
988:,
983:2
978:B
975:B
971:a
967:)
964:B
961:B
958:(
955:f
952:+
947:2
942:b
939:B
935:a
931:)
928:b
925:B
922:(
919:f
916:+
911:2
906:b
903:b
899:a
895:)
892:b
889:b
886:(
883:f
880:=
877:)
874:A
871:(
867:r
864:a
861:V
827:.
824:)
821:)
816:j
812:B
806:i
802:B
798:(
792:=
787:j
784:i
780:d
776:(
768:+
765:)
762:)
757:j
753:B
749:+
744:i
740:B
736:(
730:=
725:j
722:i
718:a
714:(
706:+
698:=
688:)
683:j
679:B
673:i
669:B
665:(
658:+
655:)
650:j
646:B
642:+
637:i
633:B
629:(
622:+
616:=
607:j
604:i
600:P
583:j
580:B
576:i
573:B
565:j
562:B
558:i
555:B
551:j
548:B
546:,
544:i
541:B
537:B
487:h
483:H
463:)
460:P
457:(
453:r
450:a
447:V
441:)
438:A
435:(
431:r
428:a
425:V
418:=
413:2
409:h
367:H
348:)
345:P
342:(
338:r
335:a
332:V
326:)
323:G
320:(
316:r
313:a
310:V
303:=
298:2
294:H
270:,
265:2
261:H
250:E
248:,
246:G
238:E
236:,
234:G
230:E
226:G
222:P
210:E
206:G
202:P
34:.
20:)
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