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We expect natural selection and genetic drift to both result in the fixation of alleles, which...

We expect natural selection and genetic drift to both result in the fixation of alleles, which seems like it should reduce average heterozygocity in populations – there should be very little observed variation. However, we observe a tremendous amount of variation in populations. What can explain this observation?

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Evolution:

In population genetics, evolution is defined as a change in the frequency of alleles (versions of a gene) in a population over time. So, evolution is any shift in allele frequencies in a population over generations – whether that shift is due to natural selection or some other evolutionary mechanism, and whether that shift makes the population better-suited for its environment or not. Both natural selection and genetic drift are mechanisms for evolution (they both change allele frequencies over time). The key distinction is that in genetic drift allele frequencies change by chance, whereas in natural selection allele frequencies change by differential reproductive success.

What is genetic drift?

Genetic drift is change in allele frequencies in a population from generation to generation that occurs due to chance events. To be more exact, genetic drift is change due to "sampling error" in selecting the alleles for the next generation from the gene pool of the current generation. Although genetic drift happens in populations of all sizes, its effects tend to be stronger in small populations.

Characteristics of genetic drift:

Population size matters

Larger populations are unlikely to change this quickly as a result of genetic drift.

Allele benefit or harm doesn't matter

Genetic drift, unlike natural selection, does not take into account an allele’s benefit (or harm) to the individual that carries it. That is, a beneficial allele may be lost, or a slightly harmful allele may become fixed, purely by chance.

A beneficial or harmful allele would be subject to selection as well as drift, but strong drift (for example, in a very small population) might still cause fixation of a harmful allele or loss of a beneficial one.

Examples of genetic drift: The bottleneck effect

The bottleneck effect is an extreme example of genetic drift that happens when the size of a population is severely reduced. Events like natural disasters (earthquakes, floods, fires) can decimate a population, killing most indviduals and leaving behind a small, random assortment of survivors.

The allele frequencies in this group may be very different from those of the population prior to the event, and some alleles may be missing entirely. The smaller population will also be more susceptible to the effects of genetic drift for generations (until its numbers return to normal), potentially causing even more alleles to be lost.

The founder effect

The founder effect is another extreme example of drift, one that occurs when a small group of individuals breaks off from a larger population to establish a colony. The new colony is isolated from the original population, and the founding individuals may not represent the full genetic diversity of the original population. That is, alleles in the founding population may be present at different frequencies than in the original population, and some alleles may be missing altogether. The founder effect is similar in concept to the bottleneck effect, but it occurs via a different mechanism (colonization rather than catastrophe).

Natural Selection

population evolution by natural selection:

  • Organisms with heritable (genetically determined) features that help them survive and reproduce in a particular environment tend to leave more offspring than their peers.
  • If this continues over generations, the heritable features that aid survival and reproduction will become more and more common in the population.
  • The population will not only evolve (change in its genetic makeup and inherited traits), but will evolve in such a way that it becomes adapted, or better-suited, to its environment.

Characteristics of Natural Selection:

Natural selection can cause microevolution

Natural selection acts on an organism’s phenotype, or observable features. Phenotype is often largely a product of genotype (the alleles, or gene versions, the organism carries). When a phenotype produced by certain alleles helps organisms survive and reproduce better than their peers, natural selection can increase the frequency of the helpful alleles from one generation to the next – that is, it can cause microevolution.

Fitness = reproductive success

The phenotypes and genotypes favoured by natural selection aren't necessarily just the ones that survive best. Instead, they're the ones with the highest overall fitness. Fitness is a measure of how well organisms survive and reproduce, with emphasis on "reproduce." Officially, fitness is defined as the number of offspring that organisms with a particular genotype or phenotype leave behind, on average, as compared to others in the population.

Mechanism of Natural Selection:

There are three basic ways that natural selection can influence distribution of phenotypes for polygenic traits in a population. To illustrate these forms of selection, let's use an imaginary beetle population, in which beetle color is controlled by many genes and varies in a spectrum from light to dark green.

Stabilizing selection. In stabilizing selection, intermediate phenotypes are more fit than extreme ones. For example, medium-green beetles might be the best camouflaged, and thus survive best, on a forest floor covered by medium-green plants. Stabilizing selection tends to narrow the curve.

Directional selection. One extreme phenotype is more fit than all the other phenotypes. For example, if the beetle population moves into a new environment with dark soil and vegetation, the dark green beetles might be better hidden and survive better than medium or light beetles. Directional selection shifts the curve towards the favorable phenotype.

Disruptive selection. Both extreme phenotypes are more fit than those in the middle. For example, if the beetles move into a new environment with patches of light-green moss and dark-green shrubs, both light and dark beetles might be better hidden (and survive better) than medium-green beetles. Diversifying selection makes multiple peaks in the curve.

Thus genetic drift and natural selection cause tremendous amount of variations in populations in the presence of favourable factors effecting them.

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