Inbreeding and its consequences, Biology tutorial

Introduction:

The broad scientific statement of inbreeding is that it is the mating of individuals more closely associated to one other than the average relationship in the population concerned. This statement is actually only valid for large populations, as with small populations inbreeding is inevitable, even by random mating. To be more specific, an inbred person is stated as someone whose parents are associated. In practice this means close direct relationship or, more generally, associated via recent common ancestors, as all members of the similar species are associated to certain extent. 

The unfavorable effects of inbreeding in animals are well recognized. The incidence of metabolic disorders, structural abnormalities and inherited disease conditions caused through injurious recessive genes, raises the following inbreeding. Performance in some characters, specifically those concerned by reproduction and survival, declines following the mating of close relatives. This is termed as inbreeding depression. Such effects are mostly due to a raise in the frequency of homozygous genotypes (AA and aa) at the expense of heterozygotes (Aa), that is caused by inbreeding. It is merely harmful, though, if the dominance is directional that signifies that the undesirable member of a pair of genes is generally recessive.

If a high proportion of such harmful genes are present in the heterozygous state (Aa) the animal is protected from their debilitating consequences through the dominance of the normal gene; however when some of the heterozygotes are substituted by homozygous recessives (aa), following inbreeding, their injurious effects become manifest. Other kinds of gene action are at times responsible for inbreeding damage, however are thought to be less significant. These are: over dominance, epistatic interaction and the overall level of heterozygosity.

Over dominance takes place if the heterozygote (A1A2) is superior in performance to either of the two homozygote (A1A1 or A2A2). In this condition, a raise in homozygosity following inbreeding as well causes inbreeding depression. 

Epistatic interaction among diverse pairs of genes takes place if one pair influences the expression of the other pair at a different locus. With one particular kind, termed as complementary epistasis, two dominants, one from each of two separate loci, are essential for normal development or metabolism. Therefore, AABB, AaBB, AABb and AaBb will be normal; however AAbb, aaBBAabb, aaBb and aabb will be defective. This condition occurs if a metabolic pathway needs two enzymes for the necessary end-product to be synthesized. As each enzyme needs a different dominant gene for its synthesis, the absence of one or both will outcome in a defective individual.

Inbreeding in a population having a mixture of the above genotypes will lead to the break-up of the favorable gene combinations, by more inferior genotypes, specifically aaBB, AAbb and aabb, being generated. 

At last, Lerner (1954) found proof that a few abnormal conditions in animals were not caused by single genes however by a drop in the general level of heterozygosity all through the whole genome. His theory of developmental homeostasis recommends that for an animal to be capable to cope with developmental accidents and environmental stress there is a minimum or obligate level of heterozygosity for normal growth. The implication being that heterozygotes in general are more versatile as they can generate a huge variety of enzymes and other proteins.

This signifies that the heterozygosity level per se, and also the effects of the genes themselves, might be a contributing factor.  The opposite of inbreeding depression is termed as heterosis or hybrid vigor and can outcome from the crossing of unrelated inbred animals or lines by various genetic backgrounds. What is lost from inbreeding is generally restored if several inbred lines are crossed arbitrarily. Deliberate inbreeding and crossing, followed by selection among lines, is at times employed by farm plants and animals to enhance the yields. Its importance in humans is that greater mobility signifies that people travel further to determine a spouse and are less probable to marry a person from the similar locality having a similar genotype. Therefore, however most of the increase in height and enhanced survival is the outcome of better nutrition and disease control, a small part might as well be due to heterosis following a change in the mating system.

The Genetic Effects of Inbreeding:

In consanguineous unions, the partners share genes inherited from one or more common ancestor and, for illustration, in first-cousin marriages the spouses are anticipated to have 12.5 percent of their genes in common. This signifies that on average their progeny will be homozygous at 6.25 percent of gene loci, that is, they will have received similar gene copies from each and every parent at such sites in their genome. If the similar mutant gene is inherited from both parents the individual will deduce the disorder, either at birth or later in life based on the nature and site of the mutation, therefore contributing to the phenomenon of inbreeding depression.

The coefficient of inbreeding (F) is a numerical approximation of the degree of inbreeding of an individual, and so for first-cousin offspring F = 0.0625. Likewise, for the progeny of uncle-niece unions F = 0.125, whereas for second-cousin offspring F = 0.0156. In most of the communities there is a long and continuous history of consanguineous unions, and therefore the cumulative level of inbreeding might be considerably higher than the value computed for a single generation. Beneath these conditions, a correction can be applied to account for the consequences of ancestral inbreeding by employing the formula:

F= 2(1/2)n (1+FA )

Here FA is the ancestor's inbreeding coefficient, n is the number of individuals in the path joining the parents of the individual, and the summation (L) is taken over each path in the pedigree which goes via a common ancestor.

The Closest Form of Inbreeding:

The closest form of inbreeding is self-fertilization that generally only takes place in monoecious animals and plants and which are hermaphrodites, example: garden peas and slugs. The equivalent condition has been experimentally generated in turkeys where a rare form of parthenogenesis takes place. Parthenogenesis (or virgin birth) is the production of viable embryos (for all time males in birds) from haploid infertile eggs through the artificial doubling of chromosome numbers. The embryos are highly homozygous. When one of such parthenogenetic males is mated back to his mother this is equivalent to self-fertilization. One generation of self-fertilization generates the similar coefficient of inbreeding (F) as three generations of full sib mating, that is, 0.5.This follows from the formula for F, which is Σ(1/2) n+1

Here, n is the number of connecting links among the two parents through a common ancestor. With selfing, both parents are the similar individual so that the number of links (n) is 0 and, thus Σ(1/2)n+1 = 0.5.

The coefficient of inbreeding:

The coefficient of inbreeding is the probability that the two genes at any locus are similar through descent (Falconer 1981), that is, that the two genes are copies of one of the genes taken through the common ancestor a few generations back. The coefficient of inbreeding, symbolized by F, is a property of an individual, however inbreeding profoundly effects the genetic composition of a population and in suitable conditions can lead to the formation of inbred strains in which all the individuals are virtually genetically similar.

The rate of inbreeding depends on the degree of relationship:

The closest relationship is that of an individual by itself, or self- fertilization. Though, the closest relationship which is generally possible having mammals is full brother x sister (termed as full-sib) mating. Continuous mating of offspring to the younger parent (that prevents repeated backcrossing to the similar individual, which would have various genetic consequences), or a single generation of parent x offspring mating is genetically equal to full-sib mating.

Other regular mating systems that lead to a high level of inbreeding comprise half-sib and cousin mating. Repeated backcrossing, state of a transgenic or a new mutation, to inbred strain rises homozygosity as quickly as self-fertilization.

Inbreeding as well arises as an outcome of restricted population size:

In a closed colony it ultimately becomes not possible to avoid the mating of associated individuals. Hence even 'outbreed' stocks maintained as a closed colony steadily become inbred at a rate which based on the size of the colony.

Full sib mating:

Full sib inbreeding of a genetically heterogeneous stock twice the net genetic variation when all the sublines are kept. Though, all the genetic variation will then be due to differences between the sublines, without genetic variation in sublines. The phenotypic variation among sublines as well rises. This is mostly due to the 'uncovering' of recessive genes and genetic drift in which alleles at a specific polymorphic locus become fixed in a homozygous state having plus or minus (with respect to the character)  alleles being fixed mainly by chance. The phenotypic variation among a set of inbred strains derived from the outbreed stock is thus substantially more than the phenotypic variation in the beginning population.

Inbreeding depression:

The Inbreeding depression is a decline in reproductive performance, capability to survive and other features related with fitness as an outcome of inbreeding. It takes place as an outcome of 'uncovering' deleterious recessive genes by making them homozygous and is an effect of the evolution of dominance of loci concerned by fitness characters. The direction of the change is towards the value of the more recessive alleles. Inbreeding depression doesn't take place for such characters where the heterozygote is intermediate among the two homozygote. 

The degree of inbreeding depression based on the prior history of the stock. A stock that has been kept as a closed population for many generations will already be partially inbred; therefore, full-sib mating might not outcome in much inbreeding depression.

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