Estimating Heritability
Since only P 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 statistical analyses required to estimate the genetic and environmental 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 twins, siblings, parents and offspring, rather than from more distantly related (and therefore less similar) subjects. The standard error for heritability estimates is improved with large sample sizes.
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 experimental control is impossible when gathering human data, relying on naturally occurring relationships and environments.
Studies of human heritability often utilise adoption study designs, often with identical twins who have been separated early in life and raised in different environments (see for example Fig. 2). 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 second and more common design is the twin study 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 not completely genetically identical, potentially resulting in an underestimation of heritability. Studies of twins also examine differences between twins and non-twin siblings, for instance to examine phenomena such as intrauterine competition (for example, twin-to-twin transfusion syndrome).
Heritability estimates are always relative to the genetic and environmental factors in the population, and are not absolute measurements of the contribution of genetic and environmental factors to a phenotype. Heritability estimates reflect the amount of variation in genotypic effects compared to variation in environmental effects.
Heritability can be made larger by diversifying the genetic background, e.g., by using only very outbred individuals (which increases VarG) and/or by minimizing environmental effects (decreasing VarE). The converse also holds. Due to such effects, different populations of a species might have different heritabilities for the same trait.
In observational studies, or because of evokative effects (where a genome evokes environments by its effect on them), G and E may covary: gene environment correlation. 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.
Heritability estimates are often misinterpreted if it is not understood that they refer to the proportion of variation between individuals in a population that is influenced by genetic factors. Heritability describes the population, not individuals within that population. For example, It is incorrect to say that since the heritability of a personality trait is about .6, that means that 60% of your personality is inherited from your parents and 40% comes from the environment.
A highly heritable trait (such as eye color) assumes environmental inputs which (though they are invariant in most populations) are required for development: for instance temperatures and atmospheres supporting life, etc.). A more useful distinction than "nature vs. nurture" is "obligate vs. facultative"—under typical environmental ranges, what traits are more "obligate" (e.g., the nose—everyone has a nose) or more "facultative" (sensitive to environmental variations, such as specific language learned during infancy). Another useful distinction is between traits that are likely to be adaptations (such as the nose) vs. those that are byproducts of adaptations (such the white color of bones), or are due to random variation (non-adaptive variation in, say, nose shape or size).
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“I am sure that in estimating every mans value either in private or public life, a pure integrity is the quality we take first into calculation, and that learning and talents are only the second.”
—Thomas Jefferson (17431826)