Micro Evolutionary

Processes-2
Mutation

Evolution requires variation. Neither selection nor drift can operate in the absence of genetic variability. Spontaneous mutation is the
primary source of all genetic variations. The more genetic variation produced by mutation, the more opportunity for evolution, and
the faster evolution can proceed. Mutation is considered as a random, non-directional and non-adaptive phenomenon. The
randomness of mutations was first demonstrated by Luria and Delbruck in 1943. The randomness of mutations is an important
concept in evolutionary biology because it is a requirement of the Darwinian view of evolution which holds that changes in the
characteristics of an organism occur by chance and are not influenced by the environment in which the organism is placed. The
Lamarckian theory of evolution on the other hand states that organisms acquire changes that enable them to adapt to their
environment. The Darwinian view requires that mutations occur at random, whereas Lamarckian evolution demands that adaptive
or directed mutations occur in response to the environment.

The occurrence of adaptive mutations was first suggested by Cairns in 1988. He found that under some circumstances, E. coli bacteria
are able to mutate their genes in a directed way that enables cells to adapt the changing environmental conditions. He worked on an
E. coli strain that carries a nonsense mutation in its lacZ gene. The presence of a nonsense mutation in lacZ means that these cells are
unable to synthesize functional b-galactosidase enzymes and so cannot use lactose as a carbon source - they are therefore lactose
auxotrophs. When the lactose auxotrophs were plated onto a minimal medium containing lactose as the only carbon source then few
lactose prototrophs appeared. This means that a second mutation occurred in the lacZ gene, reversing the effects of the nonsense
mutation and therefore allowing the b-galactosidase enzyme to be synthesized. The number of lactose prototrophs that arose was
significantly higher than that expected. Hence, it also ruled out random mutation in lacZ.

Thus, some E. coli cells underwent adaptive mutation and acquired the specific change in DNA sequence needed to withstand the
selective pressure. These experiments suggested that bacteria can program mutations according to the selective pressures. It
means that the environment can directly affect the phenotype of the organism, as suggested by Lamarck, rather than operating
through the random processes postulated by Darwin
Gene flow: A second evolutionary force that shapes these patterns is gene flow, which is the mixing of alleles from
different populations. Gene flow is mostly the outcome of either active or passive migration of individuals from one
location to the other. When the migrating individuals interbreed with the new population, they contribute their genes to
the gene pool of the local population. This establishes gene flow in a population. Gene flow plays two important roles in
evolution.
(1) It equalizes allele frequencies, and so works to erode genetic differences between populations. Natural selection can
cause two populations to become either more similar or less similar, but gene flow can only make them more similar.
(2) The second effect of gene flow is to introduce new alleles into a population from other populations where they
already exist. Here gene flow plays a role similar to that of mutation.

Gene flow results from dispersal, which is the movement of individuals and gametes. Some species disperse their genes
passively. The pollen of plants such as pines is blown many kilometers by the wind. Many plants have adaptations for
dispersing their seeds. Some animals also as the top of a plant, and spins out a thread of silk. When it feels the tug of a
breeze on the thread, the spider lets go and is carried away.
In other animals, dispersal is active. Desert locusts (Schistocerca gregaria) are grasshoppers that typically live
solitary lives. But when they are crowded and resources are scarce, changes in gene expression and hormone profiles
radically transform their behavior and morphology. They form vast swarms of hundreds of millions that fly long
distances in active search of food.

Measurement of gene flow

(1) With discrete patches, gene flow is quantified with the migration rate. This rate, symbolized by m, is the fraction of
individuals in a population that arrives from another population in each generation. If 120 individuals in a population of
size 1000 are immigrants, then the migration rate is m = 0.12. The migration rate tells us how quickly gene flow erodes
genetic differences between populations.
The difference in an allele’s frequency before and after migration
in a given population is given by the equation:

Dp = m (pm – p),

Where, pm is the allele’s frequency in the migrants and p is its

frequency in the focal population before migration. Thus, the
change in allele frequency is proportional to two quantities: the
migration rate, m, and the difference in allele frequencies between
the local population and the migrants, (pm – p).

Example:
Say that a population has an allele frequency of p = 0.25, and it
receives immigrants at a rate m = 0.1 from another population
whose allele frequency is 0.75. The above equation tells us that the
change in allele frequency is 0.05, so migration will cause the
allele frequency in the focal population to increase from p = 0.25
to 0.3. The higher the migration rate between populations, the
quicker genetic differences between them are erased.
(2) In populations that are spatially continuous, there are no
distinct populations and so the migration rate cannot be used.
Instead, we measure gene flow with the migration variance,
symbolized by sm2.
Distribution of dispersal distances of the Texas spiny lizard
(Sceloporus olivaceus, inset) in a population in Austin, Texas. The
birthplaces of individuals are centered at 0, and distribution shows the
places where they established territories and presumably reproduced.
(The original data are reported as absolute distances. This figure
assumes that half of the individuals dispersed to the left and half to the
right, which is why the distribution is exactly symmetrical.) The
variance of this distribution equals the dispersal sm2, variance, = 9800
This corresponds to an average movement of roughly sm = 99 per
generation.
Source: Evolution by Futyama