Dec.
20, 2024
Adaptive
Evolution
Adaptive Evolution
• Review:
• natural selection acts on level of the individual;
selects those with higher
overall fitness (reproductive success) compared
to rest of the population
• Natural selection favors the most adaptive variation for
a given environment
• If fit phenotypes are evolving in a stable environment,
natural selection results in stabilizing
selection
• results in overall decrease in population’s variation
• If environmental conditions change, directional
selection shifts a population’s variance
toward new (and more favorable) phenotype
• Diversifying selection results in increased variance
by selecting for two or more distinct phenotypes
Adaptive Evolution
• Sexual selection results when one sex has more reproductive success
than the other
• males and females experience different selective pressures
• often leads to distinct phenotypic differences (sexual
dimorphisms) between the two
• Example(s):
• male birds often exhibit more colorful plumage than
female birds of same species
• Important to recognize that there is no perfect organism
• Natural selection acts on existing variations in the population; does not
create anything from scratch
• although natural selection selects the fittest individuals, other
forces of evolution (genetic drift and gene flow) often introduce
deleterious alleles to population’s gene pool
• Evolution has no purpose
• it is sum of various forces that influence genetic and phenotypic
variation of a population
Adaptive Evolution
• Natural selection only acts on population’s heritable traits
• selects for beneficial alleles; increases their frequency in
the population
• selects against deleterious alleles; decreasing their
frequency
• process known as adaptive evolution
• Natural selection does not act on individual alleles, however,
but on entire organisms
• Example(s):
• individual may carry beneficial genotype (with a
resulting phenotype) that increases ability to
reproduce (fecundity)
• if same individual also carries allele that results in a
fatal childhood disease, that fecundity phenotype
will not be passed on to next generation
• individual will not live to reach reproductive
age
• Natural selection acts at level of the individual
• selects for individuals with greater contributions to the
gene pool of next generation
• known as an organism’s evolutionary (Darwinian)
fitness
• Fitness – quantifiable; measured by scientists in the field
• it is not absolute fitness of individual that counts,
Adaptive Evolution however, but rather how it compares to other
organisms in the population
• Relative fitness: concept that allows researchers to
determine which individuals are contributing additional
offspring to next generation
• how population might evolve
• Several ways selection can affect population variation:
• stabilizing selection
• directional selection
• diversifying selection
• frequency-dependent selection
• sexual selection
• As natural selection influences allele frequencies in a
population, individuals can become more/less genetically
similar
• phenotypes displayed can become more similar or
more disparate
Adaptive Evolution
(Stabilizing
Selection)
• If natural selection favors an average phenotype
(selecting against extreme variation), the population will
undergo stabilizing selection
• Example(s):
• population of mice that live in the woods
• natural selection likely to favor individuals that best
blend in with forest floor
• less likely spotted by predators
• assuming ground is consistent shade of brown, mice
whose fur is most closely matched to
that color likely to survive and
reproduce
• passing on genes for brown coat
• mice that carry alleles that make them slightly
lighter or darker will stand out against the ground
• more likely to fall victim to predation
• result of this selection: population’s genetic
variability will decrease
Adaptive Evolution (Directional Selection)
• When the environment changes, populations often undergo
directional selection
• selects for phenotypes at one end of the spectrum of existing variation
• Example(s):
• evolution of peppered moth in eighteenth- and nineteenth-century
England
• prior to Industrial Revolution, the moths were predominately light in color
• allowed them to blend in with light-colored trees and lichens in
their environment
• as soot began spewing from factories, trees became darkened
• light-colored moths became easier for predatory birds to spot
• over time, frequency of the melanic form of the moth increased
• higher survival rate in habitats affected by air pollution
• darker coloration blended with sooty trees
• Scientists can observe directional selection:
• suppose populations of rabbits that eat flowers is introduced into an
environment with flowering plants
• once flowers are eaten, plants cannot reproduce
• over time, height of flowers will shift higher so that rabbits cannot
reach them
Adaptive Evolution
(Diversifying
Selection)
• Sometimes two or more distinct phenotypes can each
have their advantages and be selected for by natural
selection
• intermediate phenotypes are (on average) less fit
• Known as diversifying selection
• Seen in many populations of animals that have multiple
male forms
• large, dominant alpha males obtain mates by brute
force
• small males can sneak in for furtive (attempt to
avoid notice) copulations with females in alpha
male’s territory
• in this case, both alpha males and “sneaking”
males will be selected for
• medium-sized males, which can’t overtake the
alpha males and are too big to sneak
Evolution
(Diversifying
Selection)
• Diversifying selection can also occur when
environmental changes favor individuals on either
end of phenotypic spectrum
• Example(s):
• population of mice living at beach where there
is light-colored sand interspersed with patches
of tall grass
• light-colored mice that blend in with sand
would be favored
• dark-colored mice also favored; can hide
in the grass
• medium-colored mice, on the other hand, would
not blend in with either grass or sand
• more likely to be eaten by predators
• Result of this type of selection: increased genetic
variability as population becomes more diverse
Adaptive Evolution
(Frequency-Dependent Selection)
• Frequency-dependent selection: favors phenotypes that are either common
(positive frequency-dependent selection) or rare (negative
frequency-dependent selection)
• Example(s):
• unique group of lizards of the Pacific Northwest
• male common side-blotched lizards come in three throat-color patterns:
• orange
• blue
• yellow
• each form has different reproductive strategy:
• orange males are strongest; can fight other males for access to their
females
• blue males are medium-sized; form strong pair bonds with their mates
• yellow males are smallest; look like females (which allows them to sneak
copulations)
• like game of rock-paper-scissors: orange beats blue, blue beats yellow, and
yellow beats orange (in competition for females)
• big, strong orange males can fight off blue males to mate with blue’s pair-
bonded females
Adaptive Evolution
(Frequency-
Dependent Selection)
• Results:
• orange males favored by natural selection when the population is
dominated by blue males
• blue males will thrive when the population is mostly yellow males
• yellow males selected for when orange males are the most
populous
• Populations of side-blotched lizards cycle in distribution of these
phenotypes:
• in one generation, orange might be predominant; yellow males
begin to rise in frequency
• once yellow males make up majority of the population, blue males
selected for
• when blue males become common, orange males once again
favored
• If selection is against the common (favoring the rare), there is negative
frequency-dependent selection
• Negative frequency-dependent selection serves to increase
population’s genetic variability
• selects for rare phenotypes
• Positive frequency-dependent selection usually decreases genetic
variability
•
Adaptive Evolution
(Sexual Selection)
• Males and females of certain species often quite different
from one another (beyond reproductive organs)
• Example(s):
• males often larger; display many elaborate colors and
adornments (peacock’s tail)
• females tend to be smaller and duller in decoration
• Such differences known as sexual dimorphisms
• arise from fact that in many populations (particularly
animal populations), there is more variability in
reproductive success of males than there is of females
• Some males (often bigger, stronger, or more decorated) get
majority of total matings; others receive none
Adaptive Evolution
(Sexual Selection)
• Occurs because males are better at fighting off other males
• or because females will choose to mate with the
bigger (more decorated) males
• In either case, this variation in reproductive success
generates a strong selection pressure among males to get
those matings
• results in evolution of bigger body size and elaborate
ornaments to get females’ attention
• Females tend to get a handful of selected matings
• more likely to select more desirable males
• Sexual dimorphism varies widely among species, of course;
some species are even sex-role reversed
• in such cases, females tend to have greater variability
in their reproductive success than males
• correspondingly selected for bigger body size
and elaborate traits (usually characteristic of
males)
Adaptive Evolution
(Sexual Selection)
• Selection pressures on males and females to obtain matings
known as sexual selection
• can result in development of secondary sexual characteristics
• do not benefit the individual’s likelihood of survival but help
maximize its reproductive success
• Sexual selection can be so strong that it selects for traits detrimental to
individual’s survival
• Example(s):
• peacock’s tail
• while beautiful and male with largest, most colorful tail is more likely
to win the female, it is not the most practical appendage
• more visible to predators; makes males slower in their
attempted escapes
• There is some evidence that the risk is why females like big tails in the first
place
• speculation: large tails carry risk
• only best males survive that risk
• Good genes hypothesis - states that males develop these
impressive ornaments to show off their efficient metabolism
(or ability to fight disease)
Adaptive • Females then choose males with most impressive traits
• signals their genetic superiority (which they will pass on
Evolution to their offspring)
(Sexual • It might be argued that females should not be so selective
Selection) • it will likely reduce their number of offspring; if better
males father more fit offspring, it may be beneficial,
however
• fewer, healthier offspring may increase chances of
survival more than many, weaker offspring
• Both handicap principle and good genes hypothesis:
• trait said to be honest signal of the males’ quality
• gives females way to find the fittest mates (males
that will pass the best genes to their offspring)
Adaptive Evolution
• Natural selection is a driving force in evolution
• can generate populations that are better adapted to
survive and successfully reproduce in their environments
• Natural selection cannot produce the perfect organism, however
• can only select on existing variation in the population
• does not create anything from scratch
• It is limited by a population’s existing genetic variability and
whatever new alleles arise through mutation and gene flow
• Natural selection also limited because it works at level of
individuals (not alleles)
• some alleles are linked due to their physical proximity in
the genome
• makes them more likely to be passed on together (linkage
disequilibrium)
Adaptive Evolution
• Any given individual may carry some
beneficial alleles and some unfavorable alleles
• It is net effect of these alleles (or organism’s
fitness) that natural selection can act
• Good alleles can be lost if carried by
individuals that also have several
overwhelmingly bad alleles
• Bad alleles can be kept if carried by
individuals that have enough good alleles to
result in an overall fitness benefit
Adaptive Evolution
• Natural selection can be constrained by relationships between different polymorphisms
• One morph may confer a higher fitness than another
• may not increase in frequency; going from less beneficial to more beneficial trait would require going through a less
beneficial phenotype
• Example(s):
• mice living at beach
• some light-colored (blend in with sand); others are dark (blend in with patches of grass)
• dark-colored mice may be more fit overall than light-colored mice
• might expect light-colored mice to be selected for darker coloration
• remember intermediate phenotype (medium-colored coat) is very bad for mice
• cannot blend in with either sand or grass
• more likely to be eaten by predators
• Result: light-colored mice would not be selected for dark coloration
• those individuals that began moving in that direction (began being selected for a darker coat) would be less fit
than those that stayed light
Adaptive Evolution
• Also important to understand that not all
evolution is adaptive
• While natural selection selects fittest
individuals and often results in a more fit
population overall, other forces of
evolution (genetic drift and gene flow)
often do the opposite
• introduce deleterious alleles to the
population’s gene pool
• Evolution has no purpose (not changing a
population into preconceived ideal)
• it is the sum of various forces and how
they influence genetic and phenotypic
variability of a population
Practice Module
Module 19.3: Adaptive Evolution
