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Genetic Load




The concept was first formulated in 1937 by JBS Haldane , independently formulated, named and applied to humans in 1950 by H. J. Muller 1, and elaborated further by Haldane in 1957.2


DEFINITION


Genetic load is the reduction in selective value for a population compared to what the population would have if all individuals had the most favored genotype.3 It is normally stated in terms of Fitness as the reduction in the mean fitness for a population compared to the maximum fitness.


MATHEMATICS

Consider a single Gene Locus with the Allele s \mathbf{A} _1 \dots \mathbf{A} _n, which have the Fitness es w_1 \dots w_n and the Allele Frequencies p_1 \dots p_n respectively. Ignoring Frequency-dependent Selection , then genetic load (L) may be calculated as:



where the i^\mathrm{th} allele is \mathbf{A}_i and has the fitness and frequency w_i and p_i respectively.

When the w_\max = 1, then (1) simplifies to

:L = 1 - \bar w. ~~~~~~~~~~(3)


CAUSES OF GENETIC LOAD

Load may be caused by selection and mutation.


Mutational load


Load caused by mutations is known as mutational load.


Selectional load


Selection occurs when the fitnesses of particular alleles are Inequal , hence selection always exerts a load.

With Directional Selection , the allele frequencies will tend towards an equilibrium position with the fittest allele reaching a frequency in Mutation-selection Balance . As mutations are rare, this is effectively fixation. Consider two alleles \mathbf{A}_1 and \mathbf{A}_2. If w_1 > w_2, then at equilibrium, p_1 \approx 1 and p_2 \approx 0, hence \bar{w} \approx w_\max, and L \approx 0.

If the mean fitness is 0, the load is equal to 1, but the population goes Extinct .


Segregational load


In contrast to directional selection, Heterozygote Advantage always exerts a load at equilibrium.


CREATIONIST CRITICISM


Some Creationists (such as Henry M. Morris ) have suggested that mutational load would increase over time and thus make populations inviable. However, they ignore the effect of selectional load acting to weed out (decrease frequency of) deleterious mutations.


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