Evolution 202

can only enhance the performance of a particular trait but an individual needs a set of beneficial mutation
to increase its overall survival and reproduction. Natural selection acts at the level of the individual within
a population; it selects for individuals with greater contributions to the gene pool of the next generation,
known as an organism’s evolutionary fitness (or Darwinian fitness).
Adaptive evolution in finches: Through natural selection, a population of finches evolved into three
separate species by adapting to several different selection pressures. Each of the three modern finches
has a beak adapted to its life history and diet

Fig 4.1. Adaptive radiation in Darwin finches

There are several ways selection can affect population variation:

1. stabilizing selection
2. directional selection
3. diversifying selection
4. frequency-dependent selection
5. sexual selection
(a) Stabilizing selection: a type of natural selection in which genetic diversity decreases as the population
stabilizes on a particular trait value
 If natural selection favours an average phenotype by selecting against extreme variation, the population
will undergo stabilizing selection. For example Robin typically laid four eggs because a larger clutch
size may result in malnourished chicks, while smaller clutches may result in no viable offspring. As a
result of this stabilizing selection, the population’s genetic variance will decrease.

(b)Directional Selection
Directional selection: a mode of natural selection in which a single phenotype is favoured, causing the
allele frequency to continuously shift in one direction
 When the environment changes, populations will often undergo directional selection, which selects for
phenotypes at one end of the spectrum of existing variation.
 POPULATION GENETICS & GENETIC DRIFT 203
 A classic example of this type of selection is the evolution of the peppered moth in eighteenth- and
nineteenth-century England. Prior to the Industrial Revolution, the moths were predominately light in
colour, which allowed them to blend in with the light-colored trees and lichens in their environment.
As soot began spewing from factories, the trees darkened and the light-colored moths became easier
for predatory birds to spot.
 Over time, the frequency of the melanic form of the moth increased because their darker coloration
provided camouflage against the sooty tree; they had a higher survival rate in habitats affected by air
pollution. Similarly, the hypothetical mouse population may evolve to take on a different coloration if
their forest floor habitat changed. The result of this type of selection is a shift in the population’s
genetic variance toward the new, fit phenotype.

(c)Diversifying (or Disruptive) Selection

Disruptive selection: A mode of natural selection in which extreme values for a trait are favoured over
intermediate values.
 Sometimes natural selection can select for two or more distinct phenotypes that each have their
advantages. In these cases, the intermediate phenotypes are often less fit than their extreme
counterparts. Known as diversifying or disruptive selection, this is seen in many populations of animals
that have multiple male mating strategies, such as lobsters. Large, dominant alpha males obtain mates
by brute force, while small males can sneak in for furtive copulations with the females in an alpha
male’s territory. In this case, both the alpha males and the “sneaking” males will be selected for, but
medium-sized males, which cannot overtake the alpha males and are too big to sneak in for copulations,
are selected against.
 Diversifying selection can also occur when environmental changes favour individuals on either end of
the phenotypic spectrum. Let’s take an example of population of gray and Himalayan rabbits (grey
and white), Himalayan rabbits are better able to blend with the rocky environment than white rabbits,
resulting in diversifying selection. The result of this type of selection is increased genetic variance
as the population becomes more diverse.

The Lederberg experiment

In 1952, Esther and Joshua Lederberg performed an experiment that helped to show that many mutations
are random, not directed.
Here is the experimental set-up for the Lederberg experiment. All you really need to know in terms of
 Evolution 204
background information is that bacteria grow into isolated colonies on plates, and that you can reproduce
the colonies from an original plate to new plates by “stamping” the original plate with a cloth and then
stamping empty plates with the same cloth. Bacteria from each colony are picked up on the cloth and then
deposited on the new plates by the cloth.
The hypothesis for the experiment is that antibiotic resistant strains of bacteria surviving an application of
antibiotics had the resistance before their exposure to the antibiotics, not as a result of the exposure.
1. Bacteria are spread out on a plate, called the “original plate.”
2. They are allowed to grow into several different colonies.
3. This layout of colonies is stamped from the original plate onto a new plate that contains the antibiotic
penicillin.
4. Colonies X and Y on the stamped plate survive. They must carry a mutation for penicillin resistance.
5. The Lederberg’s set out to answer the question; did the colonies on the new plate evolve antibiotic
resistance because they were exposed to penicillin? The answer is no:When the original plate is
washed with penicillin, the same colonies (those in position X and Y) live — even though these
colonies on the original plate have never encountered penicillin before.
So the penicillin-resistant bacteria were there in the population before they encountered penicillin.
They did not evolve resistance in response to exposure to the antibiotic.

Fig. 4.2. Esther and Joshua Lederberg experiment

Thus, different mutations exist in population and are selected for expression when the environment is
appropriate.
Genetic drift or Sewall-Wright effect
In each generation, some individuals may, just by chance, leave behind a few more descendents (and
genes, of course!) than other individuals. The genes of the next generation will be the genes of the “lucky”
individuals, not necessarily the healthier or “better” individuals. That, in a nutshell, is genetic drift. It
happens to all populations — there’s no avoiding the vagaries of chance.
Genetic drift affects the genetic makeup of the population but, unlike natural selection, through an entirely
random process. So although genetic drift is a mechanism of evolution, it doesn’t work to
produce adaptations.
 POPULATION GENETICS & GENETIC DRIFT 205
Bottlenecks and founder effects
Genetic drift can cause big losses of genetic variation for small populations.
Population bottlenecks occur when a population’s size is reduced for at least one generation. Because
genetic drift acts more quickly to reduce genetic variation in small populations, undergoing a bottleneck
can reduce a population’s genetic variation by a lot, even if the bottleneck doesn’t last for very many
generations. This is illustrated by the bags of marbles shown below, where, in generation 2; an unusually
small draw creates a bottleneck.
Reduced genetic variation means that the population may not be able to adapt to new selection pressures,
such as climatic change or a shift in available resources, because the genetic variation that selection would
act on may have already drifted out of the population.

Fig 4.2. Population bottlenecks occur when a population’s size is reduced for at least one generation. Because
genetic drift acts more quickly to reduce genetic variation in small populations, undergoing a bottleneck can
reduce a population’s genetic variation by a lot, even if the bottleneck doesn’t last for many generations. This is
illustrated by the bags of marbles shown above here red marbles completely lost from the sample in the final
generation.
An example of a bottleneck
Northern elephant seals have reduced genetic variation probably because of a population bottleneck
humans inflicted on them in the 1890s. Hunting reduced their population size to as few as 20 individuals
at the end of the 19th century. Their population has since rebounded to over 30,000 — but their genes
still carry the marks of this bottleneck: they have much less genetic variation than a population of southern
elephant seals that was not so intensely hunted.
Founder effect
A founder effect occurs when a new colony is started by a few members of the original population. This
small population size means that the colony may have:
Reduced genetic variation from the original population.
A non-random sample of the genes in the original population.

Fig. 4.3. Founders effect occurs when a new colony is started by a few members of the original population.

For example, the African population of Dutch settlers in South Africa is descended mainly from a few
colonists. Today, the Afrikaner population has an unusually high frequency of the gene that causes
Huntington’s disease, because those original Dutch colonists just happened to carry that gene with unusually
 Evolution 206
high frequency. This effect is easy to recognize in genetic diseases, but of course, the frequencies of all
sorts of genes are affected by founder events.
Another example; when humans take a few individuals of some organism and try to establish a population
of them in some other country where they do not exist (founder effect) or when humans kill a lot of
individual of some population (ex. Tigers and leopards in India) so that only few individuals remain
(bottleneck effect).
Example-1: Genetic variability in the natural populations is generated by [B.H.U.-2015]
(a) Genetic drift (b) Sewall-Wright effect
(c) Mutation (d) Selection
Ans. (c) Mutations
Soln. It is a permanent alternation of the nucleotide sequence of the Genome of an organism and it causes the
genetic variability in natural population.
Example-2: Due to founder effect in a population, the degree of genetic variability:
(a) Is similar to original population (b) Remains constant [B.H.U.-2016]
(c) Is increased (d) is reduced
Soln. In founder effect, only few individual seek & adapt in new place. And upcoming generations of this
species at this place show genetic composition of this group of individual. But, this group is not representing
the actual or original gene pool of their parent population. So, their genetic diversity of this founder’s
population is somewhat less then original population.