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Population variation – Selection
Assignment
Subject : Crop Evolution GPB821
Presented by: Mr. Indranil Bhattacharjee
Student I.D. No.: 17PHGPB102
Presented to : Dr. G.M. Lal
Sam Higginbottom University of Agriculture, Technology &
Sciences
Allahabad-211007
Overview: The Smallest Unit of Evolution
• One common misconception is that organisms
evolve during their lifetimes
• Natural selection acts on individuals, but only
populations evolve
• Consider, for example, a population of medium
ground finches on Daphne Major Island
– During a drought, large-beaked birds were more likely
to crack large seeds and survive
– The finch population evolved by natural selection
Figure 21.2
1978
(after
drought)
10
1976
(similar to the
prior 3 years)
Averagebeakdepth(mm)
9
8
0
• Microevolution is a change in allele frequencies
in a population over generations
• Three mechanisms cause allele frequency change
– Natural selection
– Genetic drift
– Gene flow
• Only natural selection causes adaptive evolution
• Variation in heritable traits is a prerequisite for
evolution
• Mendel’s work on pea plants provided evidence
of discrete heritable units (genes)
Concept 21.1: Genetic variation makes
evolution possible
Genetic Variation
• Phenotypic variation often reflects genetic
variation
• Genetic variation among individuals is caused by
differences in genes or other DNA sequences
• Some phenotypic differences are due to
differences in a single gene and can be classified
on an “either-or” basis
• Other phenotypic differences are due to the
influence of many genes and vary in gradations
along a continuum
Figure 21.3
• Genetic variation can be measured at the whole
gene level as gene variability
• Gene variability can be quantified as the average
percent of loci that are heterozygous
• Genetic variation can be measured at the
molecular level of DNA as nucleotide variability
• Nucleotide variation rarely results in phenotypic
variation
• Phenotype is the product of inherited genotype
and environmental influences
• Natural selection can only act on phenotypic
variation that has a genetic component
Figure 21.5
(a) Caterpillars raised on a diet of
oak flowers
(b) Caterpillars raised on a diet of
oak leaves
Sources of Genetic Variation
• New genes and alleles can arise by mutation or
gene duplication
Formation of New Alleles
• A mutation is a change in the nucleotide
sequence of DNA
• Only mutations in cells that produce gametes can
be passed to offspring
• A “point mutation” is a change in one base in a
gene
• The effects of point mutations can vary
– Mutations in noncoding regions of DNA are often
harmless
– Mutations to genes can be neutral because of
redundancy in the genetic code
• The effects of point mutations can vary
– Mutations that alter the phenotype are often harmful
– Mutations that result in a change in protein
production can sometimes be beneficial
Altering Gene Number or Position
• Chromosomal mutations that delete, disrupt, or
rearrange many loci are typically harmful
• Duplication of small pieces of DNA increases
genome size and is usually less harmful
• Duplicated genes can take on new functions by
further mutation
• An ancestral odor-detecting gene has been
duplicated many times: Humans have 350
functional copies of the gene; mice have 1,000
Rapid Reproduction
• Mutation rates are low in animals and plants
• The average is about one mutation in every
100,000 genes per generation
• Mutation rates are often lower in prokaryotes and
higher in viruses
• Short generation times allow mutations to
accumulate rapidly in prokaryotes and viruses
Sexual Reproduction
• In organisms that reproduce sexually, most
genetic variation results from recombination of
alleles
• Sexual reproduction can shuffle existing alleles
into new combinations through three
mechanisms: crossing over, independent
assortment, and fertilization
Concept 21.2: The Hardy-Weinberg equation can be
used to test whether a population is evolving
• The first step in testing whether evolution is
occurring in a population is to clarify what we
mean by a population
Gene Pools and Allele Frequencies
• A population is a localized group of individuals
capable of interbreeding and producing fertile
offspring
• A gene pool consists of all the alleles for all loci in
a population
• An allele for a particular locus is fixed if all
individuals in a population are homozygous for
the same allele
Figure 21.6
Porcupine herd
Beaufort Sea
Fortymile herd
Porcupine
herd range
Fortymile
herd range
MAP
AREA
ALASKA
CANADA
• The frequency of an allele in a population can be
calculated
– For diploid organisms, the total number of alleles at a
locus is the total number of individuals times 2
– The total number of dominant alleles at a locus is 2
alleles for each homozygous dominant individual plus
1 allele for each heterozygous individual; the same
logic applies for recessive alleles
• By convention, if there are 2 alleles at a locus, p
and q are used to represent their frequencies
• The frequency of all alleles in a population will
add up to 1
– For example, p  q  1
Figure 21.UN01
CR
CR
CR
CW
CW
CW
• For example, consider a population of wildflowers
that is incompletely dominant for color
– 320 red flowers (CRCR)
– 160 pink flowers (CRCW)
– 20 white flowers (CWCW)
• Calculate the number of copies of each allele
– CR  (320  2)  160  800
– CW  (20  2)  160  200
• To calculate the frequency of each allele
– p  freq CR  800 / (800  200)  0.8 (80%)
– q  1  p  0.2 (20%)
• The sum of alleles is always 1
– 0.8  0.2  1
The Hardy-Weinberg Principle
• The Hardy-Weinberg principle describes a
population that is not evolving
• If a population does not meet the criteria of the
Hardy-Weinberg principle, it can be concluded
that the population is evolving
• The Hardy-Weinberg principle states that
frequencies of alleles and genotypes in a
population remain constant from generation to
generation
• In a given population where gametes contribute
to the next generation randomly, allele
frequencies will not change
• Mendelian inheritance preserves genetic
variation in a population
Hardy-Weinberg Equilibrium
• Hardy-Weinberg equilibrium describes the
constant frequency of alleles in such a gene pool
• Consider, for example, the same population of
500 wildflowers and 1,000 alleles where
– p  freq CR  0.8
– q  freq CW  0.2
Figure 21.7
Frequencies of alleles
Gametes produced
p  frequency of CR
allele
q  frequency of CW
allele
Alleles in the population
Each egg: Each sperm:
 0.8
 0.2
80%
chance
80%
chance
20%
chance
20%
chance
• The frequency of genotypes can be calculated
– CRCR  p2  (0.8)2  0.64
– CRCW  2pq  2(0.8)(0.2)  0.32
– CWCW  q2  (0.2)2  0.04
• The frequency of genotypes can be confirmed
using a Punnett square
Figure 21.8
Sperm
Eggs
80% CR
(p  0.8) 20% CW
(q  0.2)
p  0.8 q  0.2CR
CR
CW
CW
p  0.8
q  0.2
0.64 (p2
)
CR
CR
0.16 (pq)
CR
CW
0.16 (qp)
CR
CW
0.04 (q2
)
CW
CW
Gametes of this generation:
64% CR
CR
, 32% CR
CW
, and 4% CW
CW
64% CR
(from CR
CR
plants)
16% CR
(from CR
CW
plants)
4% CW
(from CW
CW
plants)
16% CW
(from CR
CW
plants)
80% CR
 0.8  p
20% CW
 0.2  q




64% CR
CR
, 32% CR
CW
, and 4% CW
CW
plants
With random mating, these gametes will result in the same
mix of genotypes in the next generation:
Figure 21.8a
Sperm
Eggs
80% CR
(p  0.8) 20% CW
(q  0.2)
p  0.8 q  0.2CR
CR
CW
CW
p  0.8
q  0.2
0.64 (p2
)
CR
CR
0.16 (pq)
CR
CW
0.16 (qp)
CR
CW
0.04 (q2
)
CW
CW
Figure 21.8b
Gametes of this generation:
64% CR
CR
, 32% CR
CW
, and 4% CW
CW
64% CR
(from CR
CR
plants)
16% CR
(from CR
CW
plants)
4% CW
(from CW
CW
plants)
16% CW
(from CR
CW
plants)
80% CR
 0.8  p
20% CW
 0.2  q




64% CR
CR
, 32% CR
CW
, and 4% CW
CW
plants
With random mating, these gametes will result in the same
mix of genotypes in the next generation:
• If p and q represent the relative frequencies of
the only two possible alleles in a population at a
particular locus, then
– p2  2pq  q2  1
where p2 and q2 represent the frequencies of the
homozygous genotypes and 2pq represents the
frequency of the heterozygous genotype
Figure 21.UN02
Conditions for Hardy-Weinberg Equilibrium
• The Hardy-Weinberg theorem describes a
hypothetical population that is not evolving
• In real populations, allele and genotype
frequencies do change over time
• The five conditions for nonevolving populations
are rarely met in nature
1. No mutations
2. Random mating
3. No natural selection
4. Extremely large population size
5. No gene flow
• Natural populations can evolve at some loci while
being in Hardy-Weinberg equilibrium at other loci
• Some populations evolve slowly enough that
evolution cannot be detected
Applying the Hardy-Weinberg Principle
• We can assume the locus that causes
phenylketonuria (PKU) is in Hardy-Weinberg
equilibrium given that
1. The PKU gene mutation rate is low
2. Mate selection is random with respect to whether or
not an individual is a carrier for the PKU allele
3. Natural selection can only act on rare homozygous
individuals who do not follow dietary restrictions
4. The population is large
5. Migration has no effect, as many other populations
have similar allele frequencies
• The occurrence of PKU is 1 per 10,000 births
– q2  0.0001
– q  0.01
• The frequency of normal alleles is
– p  1 – q  1 – 0.01  0.99
• The frequency of carriers is
– 2pq  2  0.99  0.01  0.0198
– or approximately 2% of the U.S. population
• Three major factors alter allele frequencies and
bring about most evolutionary change
– Natural selection
– Genetic drift
– Gene flow
Concept 21.3: Natural selection, genetic drift, and
gene flow can alter allele frequencies in a population
Natural Selection
• Differential success in reproduction results in
certain alleles being passed to the next generation
in greater proportions
• For example, an allele that confers resistance to
DDT increased in frequency after DDT was used
widely in agriculture
Genetic Drift
• The smaller a sample, the more likely it is that
chance alone will cause deviation from a predicted
result
• Genetic drift describes how allele frequencies
fluctuate unpredictably from one generation to the
next
• Genetic drift tends to reduce genetic variation
through losses of alleles, especially in small
populations
Figure 21.9-1
CW
CW
CR
CR
CR
CW
CR
CR
CR
CR
CR
CR
CR
CR
CR
CW
CR
CW
CR
CW
p (frequency of CR
)  0.7
q (frequency of CW
)  0.3
Generation 1
Figure 21.9-2
CW
CW
CR
CR
CR
CW
CR
CR
CR
CR
CR
CR
CR
CR
CR
CW
CR
CW
CR
CW
CW
CW
CR
CR
CR
CW
CR
CR
CR
CR
CR
CW
CR
CW
CR
CW
CW
CW
CW
CW
5 plants
leave
offspring
p (frequency of CR
)  0.7
q (frequency of CW
)  0.3
p  0.5
q  0.5
Generation 2Generation 1
Figure 21.9-3
CW
CW
CR
CR
CR
CW
CR
CR
CR
CR
CR
CR
CR
CR
CR
CW
CR
CW
CR
CW
CW
CW
CR
CR
CR
CW
CR
CR
CR
CR
CR
CR
CR
CR
CR
CW
CR
CW
CR
CW
CW
CW
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CR
CW
CW
5 plants
leave
offspring
2 plants
leave
offspring
p (frequency of CR
)  0.7
q (frequency of CW
)  0.3
p  0.5
q  0.5
p  1.0
q  0.0
Generation 2 Generation 3Generation 1
The Founder Effect
• The founder effect occurs when a few individuals
become isolated from a larger population
• Allele frequencies in the small founder
population can be different from those in the
larger parent population due to chance
The Bottleneck Effect
• The bottleneck effect can result from a drastic
reduction in population size due to a sudden
environmental change
• By chance, the resulting gene pool may no longer
be reflective of the original population’s gene pool
• If the population remains small, it may be further
affected by genetic drift
Figure 21.10
Original
population
Surviving
population
Bottlenecking
event
(a) By chance, blue marbles are overrepresented in
the surviving population.
(b) Florida panther (Puma concolor coryi)
Figure 21.10a-1
Original
population
(a) By chance, blue marbles are overrepresented in the
surviving population.
Figure 21.10a-2
Original
population
Bottlenecking
event
(a) By chance, blue marbles are overrepresented in the
surviving population.
Figure 21.10a-3
Original
population
Surviving
population
Bottlenecking
event
(a) By chance, blue marbles are overrepresented in the
surviving population.
Figure 21.10b
(b) Florida panther (Puma concolor coryi)
• Understanding the bottleneck effect can
increase understanding of how human activity
affects other species
Case Study: Impact of Genetic Drift on
the Greater Prairie Chicken
• Loss of prairie habitat caused a severe reduction
in the population of greater prairie chickens in
Illinois
• The surviving birds had low levels of genetic
variation, and only 50% of their eggs hatched
Figure 21.11
Pre-bottleneck
(Illinois, 1820)
Post-bottleneck
(Illinois, 1993)
Range
of greater
prairie
chicken
Illinois
1930–1960s
1993
Greater prairie chicken
Kansas, 1998
(no bottleneck)
Nebraska, 1998
(no bottleneck)
1,000–25,000
50
75,000–
200,000
5.2
3.7
Location Population
size
750,000
Number
of alleles
per locus
Percentage
of eggs
hatched
93
50
5.8
5.8
99
96
(a)
(b)
Figure 21.11a
Pre-bottleneck
(Illinois, 1820)
Post-bottleneck
(Illinois, 1993)
Range
of greater
prairie
chicken
Greater prairie chicken
(a)
• Researchers used DNA from museum specimens
to compare genetic variation in the population
before and after the bottleneck
• The results showed a loss of alleles at several loci
• Researchers introduced greater prairie chickens
from populations in other states and were
successful in introducing new alleles and
increasing the egg hatch rate to 90%
Effects of Genetic Drift: A Summary
1. Genetic drift is significant in small populations
2. Genetic drift can cause allele frequencies to
change at random
3. Genetic drift can lead to a loss of genetic
variation within populations
4. Genetic drift can cause harmful alleles to become
fixed
Gene Flow
• Gene flow consists of the movement of alleles
among populations
• Alleles can be transferred through the movement
of fertile individuals or gametes (for example,
pollen)
• Gene flow tends to reduce genetic variation
among populations over time
• Gene flow can decrease the fitness of a
population
• Consider, for example, the great tit (Parus major)
on the Dutch island of Vlieland
– Immigration of birds from the mainland introduces
alleles that decrease fitness in island populations
– Natural selection reduces the frequency of these
alleles in the eastern population where immigration
from the mainland is low
– In the central population, high immigration from the
mainland overwhelms the effects of selection
Figure 21.12
Survivalrate(%)
Central
population
Vlieland,
the Netherlands
Eastern
population
NORTH SEA
2 km
Population in which the
surviving females
eventually bred
Females born in
central population
Parus major
Central
Eastern
Females born in
eastern population
60
50
40
30
20
10
0
• Gene flow can increase the fitness of a
population
• Consider, for example, the spread of alleles for
resistance to insecticides
– Insecticides have been used to target mosquitoes that
carry West Nile virus and other diseases
– Alleles have evolved in some populations that confer
insecticide resistance to these mosquitoes
– The flow of insecticide resistance alleles into a
population can cause an increase in fitness
• Gene flow is an important agent of evolutionary
change in modern human populations
• Evolution by natural selection involves both
chance and “sorting”
– New genetic variations arise by chance
– Beneficial alleles are “sorted” and favored by natural
selection
• Only natural selection consistently results in
adaptive evolution, an increase in the frequency
of alleles that improve fitness
Concept 21.4: Natural selection is the only
mechanism that consistently causes adaptive
evolution
Natural Selection: A Closer Look
• Natural selection brings about adaptive evolution
by acting on an organism’s phenotype
Relative Fitness
• The phrases “struggle for existence” and “survival
of the fittest” are misleading as they imply direct
competition among individuals
• Reproductive success is generally more subtle and
depends on many factors
• Relative fitness is the contribution an individual
makes to the gene pool of the next generation,
relative to the contributions of other individuals
• Selection indirectly favors certain genotypes by
acting directly on phenotypes
Directional, Disruptive, and Stabilizing
Selection
• There are three modes of natural selection
– Directional selection favors individuals at one end of
the phenotypic range
– Disruptive selection favors individuals at both
extremes of the phenotypic range
– Stabilizing selection favors intermediate variants and
acts against extreme phenotypes
Figure 21.13
Original
population
Evolved
population
Original
population
Frequencyof
individuals
Phenotypes (fur color)
(a) Directional selection (b) Disruptive selection (c) Stabilizing selection
The Key Role of Natural Selection in
Adaptive Evolution
• Striking adaptations have arisen by natural
selection
– For example, certain octopuses can change color
rapidly for camouflage
– For example, the jaws of snakes allow them to
swallow prey larger than their heads
Figure 21.14
Bones shown in
green are movable.
Ligament
• Natural selection increases the frequencies of
alleles that enhance survival and reproduction
• Adaptive evolution occurs as the match between
an organism and its environment increases
• Because the environment can change, adaptive
evolution is a continuous, dynamic process
• Genetic drift and gene flow do not consistently
lead to adaptive evolution, as they can increase
or decrease the match between an organism and
its environment
Sexual Selection
• Sexual selection is natural selection for mating
success
• It can result in sexual dimorphism, marked
differences between the sexes in secondary sexual
characteristics
Figure 21.15
• Intrasexual selection is competition among
individuals of one sex (often males) for mates of
the opposite sex
• Intersexual selection, often called mate choice,
occurs when individuals of one sex (usually
females) are choosy in selecting their mates
• Male showiness due to mate choice can increase
a male’s chances of attracting a female, while
decreasing his chances of survival
• How do female preferences evolve?
• The “good genes” hypothesis suggests that if a
trait is related to male genetic quality or health,
both the male trait and female preference for that
trait should increase in frequency
Figure 21.16
SC male gray
tree frog
LC male gray
tree frog
Female gray
tree frog
Recording of SC
male’s call
Recording of LC
male’s call
Offspring of
SC father
SC sperm  Eggs  LC sperm
Offspring of
LC father
Survival and growth of these half-sibling offspring compared
Experiment
Results
Figure 21.16a
SC male gray
tree frog
LC male gray
tree frog
Female gray
tree frog
Recording of SC
male’s call
Recording of LC
male’s call
Offspring of
SC father
SC sperm  Eggs  LC sperm
Offspring of
LC father
Survival and growth of these half-sibling offspring compared
Experiment
The Preservation of Genetic Variation
• Neutral variation is genetic variation that does
not confer a selective advantage or disadvantage
• Various mechanisms help to preserve genetic
variation in a population
Diploidy
• Diploidy maintains genetic variation in the form
of hidden recessive alleles
• Heterozygotes can carry recessive alleles that are
hidden from the effects of selection
Balancing Selection
• Balancing selection occurs when natural
selection maintains stable frequencies of two or
more phenotypic forms in a population
• Balancing selection includes
– Heterozygote advantage
– Frequency-dependent selection
• Heterozygote advantage occurs when
heterozygotes have a higher fitness than do both
homozygotes
• Natural selection will tend to maintain two or
more alleles at that locus
• For example, the sickle-cell allele causes
deleterious mutations in hemoglobin but also
confers malaria resistance
Figure 21.17
Distribution of
malaria caused by
Plasmodium falciparum
(a parasitic unicellular eukaryote)
Key
Frequencies of the
sickle-cell allele
10.0–12.5%
12.5%
7.5–10.0%
5.0–7.5%
2.5–5.0%
0–2.5%
• Frequency-dependent selection occurs when the
fitness of a phenotype declines if it becomes too
common in the population
• Selection can favor whichever phenotype is less
common in a population
• For example, frequency-dependent selection
selects for approximately equal numbers of
“right-mouthed” and “left-mouthed” scale-
eating fish
Figure 21.18
“Left-mouthed”
P. microlepis
“Right-mouthed”
P. microlepis
Sample year
Frequencyof
“left-mouthed”individuals
1981 ’83 ’85 ’87 ’89
0.5
0
1.0
KEY CONCEPT
A population shares a common gene pool.
Genetic variation in a population increases
the chance that some individuals will survive.
• Genetic variation leads to phenotypic variation.
• Phenotypic variation is necessary for natural selection.
• Genetic variation is stored in a population’s gene pool.
– made up of all alleles in a population
– allele combinations form when organisms have offspring
• Allele frequencies measure genetic variation.
– measures how common allele is in population
– can be calculated for each allele in gene pool
Genetic variation comes from several
sources.
• Mutation is a random change in the DNA of a gene.
• Recombination forms new combinations of alleles.
– can form new allele
– can be passed on to
offspring if in
reproductive cells
– usually occurs during meiosis
– parents’ alleles
arranged in new
ways in gametes
Genetic variation comes from several
sources.
• Hybridization is the crossing of two different species.
– occurs when individuals can’t find mate of own species
– topic of current scientific research
KEY CONCEPT
Populations, not individuals, evolve.
Natural selection acts on distributions of
traits.
• A normal distribution graphs as a bell-shaped curve.
• Traits not undergoing natural
selection have a normal distribution.
– highest frequency near
mean value
– frequencies decrease
toward each extreme
value
Natural selection can change the
distribution of a trait in one of three
ways.
• Microevolution is evolution within a population.
– observable change in the allele frequencies
– can result from natural selection
• Natural selection can take one of three
paths.
– Directional selection favors phenotypes at one extreme.
– Stabilizing selection favors the intermediate
phenotype.
• Natural selection can take one of three
paths.
• Natural selection can take one of
three paths.
– Disruptive selection favors both extreme
phenotypes.
KEY CONCEPT
Natural selection is not the only mechanism through which populations
evolve.
Gene flow is the movement of alleles
between populations.
• Gene flow occurs when
individuals join new
populations and
reproduce.
• Gene flow keeps
neighboring populations
similar.
• Low gene flow increases
the chance that two
populations will evolve
into different species.
bald eagle migration
Genetic drift is a change in allele
frequencies due to chance.
• Genetic drift causes a loss of genetic diversity.
• It is most common in small populations.
• A population bottleneck can lead to genetic drift.
– It occurs when an event
drastically reduces
population size.
– The bottleneck effect is
genetic drift that occurs
after a bottleneck event.
• The founding of a small population can lead to genetic drift.
– It occurs when a few individuals start a new population.
– The founder effect is genetic drift that occurs after start of new population.
• Genetic drift has negative effects on a population.
– less likely to have some individuals that can adapt
– harmful alleles can become more common due to chance
Sexual selection occurs when certain
traits increase mating success.
• Sexual selection occurs
due to higher cost of
reproduction for females.
– males produce many
sperm continuously
– females are more limited
in potential offspring each
cycle
• There are two types of sexual selection.
– intrasexual selection: competition among males
– intersexual selection: males display certain traits to females
KEY CONCEPT
New species can arise when populations are isolated.
The isolation of populations can lead to
speciation.
• Populations become isolated when there is no gene flow.
– Isolated populations adapt to their own environments.
– Genetic differences can add up over generations.
• Reproductive isolation can occur between isolated
populations.
– members of different
populations cannot mate
successfully
– final step to becoming
separate species
• Speciation is the rise of two or more species from one existing species.
Populations can become isolated in
several ways.
• Behavioral barriers can cause isolation.
– called behavioral isolation
– includes differences in courtship or mating behaviors
• Geographic barriers can cause isolation.
– called geographic isolation
– physical barriers divide population
• Temporal barriers can cause isolation.
– called temporal isolation
– timing of reproductive periods prevents mating
KEY CONCEPT
Evolution occurs in patterns.
Evolution through natural selection is not
random.
• Natural selection can have direction.
• The effects of natural selection add up over time.
• Convergent evolution describes evolution
toward similar traits in unrelated species.
• Divergent evolution describes evolution toward different
traits in closely related species.
How do convergent and divergent
evolution illustrate the directional
nature of natural selection?
ancestor
red foxkit fox
Species can shape each other over time.
• Two or more species can evolve together through
coevolution.
– evolutionary paths become connected
– species evolve in response to changes in each other
• Coevolution can occur in beneficial relationships.
• Coevolution can occur in competitive relationships,
sometimes called evolutionary.
Species can become extinct.
• Extinction is the elimination of a species from Earth.
• Background extinctions occur continuously at a very low
rate.
– occur at roughly the same rate as
speciation
– usually affects a few species in a
small area
– caused by local changes in
environment
• Background extinctions occur continuously at a very low
rate.
– occur at roughly the same rate as speciation
– usually affects a few species in a small area
– caused by local changes in environment
• Mass extinctions are rare but much more intense.
– destroy many species at global level
– thought to be caused by catastrophic events
– at least five mass extinctions in last 600 million years
Speciation often occurs in patterns.
• A pattern of punctuated equilibrium exists in the
fossil record.
– theory proposed by Eldredge and Gould in 1972
– episodes of speciation occur suddenly in geologic
time
– followed by long periods of little evolutionary change
– revised Darwin’s idea that species arose through
gradual transformations
• Many species evolve from one species during adaptive
radiation.
– ancestral species diversifies into many descendent species
– descendent species
usually adapted to
wide range of
environments
Moral of this story : Survival of the fittest…….
References
James F. Hancock 2012 Plant Evolution and the Origin of
Crop Species 3rd eds CABI Publishing
John D. Thompson 2005 Plant Evolution in the
Mediterranean Oxford University Press
Miller AJ. 2007. Crop Plants: Evolution. John Wiley & Sons.
Peter R. Bell, Alan R. Hemsley 1992 Green Plants Their
Origin and Diversity Cambridge University Press

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05 population variation and selection

  • 1. Population variation – Selection Assignment Subject : Crop Evolution GPB821 Presented by: Mr. Indranil Bhattacharjee Student I.D. No.: 17PHGPB102 Presented to : Dr. G.M. Lal Sam Higginbottom University of Agriculture, Technology & Sciences Allahabad-211007
  • 2. Overview: The Smallest Unit of Evolution • One common misconception is that organisms evolve during their lifetimes • Natural selection acts on individuals, but only populations evolve • Consider, for example, a population of medium ground finches on Daphne Major Island – During a drought, large-beaked birds were more likely to crack large seeds and survive – The finch population evolved by natural selection
  • 3. Figure 21.2 1978 (after drought) 10 1976 (similar to the prior 3 years) Averagebeakdepth(mm) 9 8 0
  • 4. • Microevolution is a change in allele frequencies in a population over generations • Three mechanisms cause allele frequency change – Natural selection – Genetic drift – Gene flow • Only natural selection causes adaptive evolution
  • 5. • Variation in heritable traits is a prerequisite for evolution • Mendel’s work on pea plants provided evidence of discrete heritable units (genes) Concept 21.1: Genetic variation makes evolution possible
  • 6. Genetic Variation • Phenotypic variation often reflects genetic variation • Genetic variation among individuals is caused by differences in genes or other DNA sequences • Some phenotypic differences are due to differences in a single gene and can be classified on an “either-or” basis • Other phenotypic differences are due to the influence of many genes and vary in gradations along a continuum
  • 8. • Genetic variation can be measured at the whole gene level as gene variability • Gene variability can be quantified as the average percent of loci that are heterozygous
  • 9. • Genetic variation can be measured at the molecular level of DNA as nucleotide variability • Nucleotide variation rarely results in phenotypic variation
  • 10. • Phenotype is the product of inherited genotype and environmental influences • Natural selection can only act on phenotypic variation that has a genetic component
  • 11. Figure 21.5 (a) Caterpillars raised on a diet of oak flowers (b) Caterpillars raised on a diet of oak leaves
  • 12. Sources of Genetic Variation • New genes and alleles can arise by mutation or gene duplication
  • 13. Formation of New Alleles • A mutation is a change in the nucleotide sequence of DNA • Only mutations in cells that produce gametes can be passed to offspring • A “point mutation” is a change in one base in a gene
  • 14. • The effects of point mutations can vary – Mutations in noncoding regions of DNA are often harmless – Mutations to genes can be neutral because of redundancy in the genetic code
  • 15. • The effects of point mutations can vary – Mutations that alter the phenotype are often harmful – Mutations that result in a change in protein production can sometimes be beneficial
  • 16. Altering Gene Number or Position • Chromosomal mutations that delete, disrupt, or rearrange many loci are typically harmful • Duplication of small pieces of DNA increases genome size and is usually less harmful • Duplicated genes can take on new functions by further mutation • An ancestral odor-detecting gene has been duplicated many times: Humans have 350 functional copies of the gene; mice have 1,000
  • 17. Rapid Reproduction • Mutation rates are low in animals and plants • The average is about one mutation in every 100,000 genes per generation • Mutation rates are often lower in prokaryotes and higher in viruses • Short generation times allow mutations to accumulate rapidly in prokaryotes and viruses
  • 18. Sexual Reproduction • In organisms that reproduce sexually, most genetic variation results from recombination of alleles • Sexual reproduction can shuffle existing alleles into new combinations through three mechanisms: crossing over, independent assortment, and fertilization
  • 19. Concept 21.2: The Hardy-Weinberg equation can be used to test whether a population is evolving • The first step in testing whether evolution is occurring in a population is to clarify what we mean by a population
  • 20. Gene Pools and Allele Frequencies • A population is a localized group of individuals capable of interbreeding and producing fertile offspring • A gene pool consists of all the alleles for all loci in a population • An allele for a particular locus is fixed if all individuals in a population are homozygous for the same allele
  • 21. Figure 21.6 Porcupine herd Beaufort Sea Fortymile herd Porcupine herd range Fortymile herd range MAP AREA ALASKA CANADA
  • 22. • The frequency of an allele in a population can be calculated – For diploid organisms, the total number of alleles at a locus is the total number of individuals times 2 – The total number of dominant alleles at a locus is 2 alleles for each homozygous dominant individual plus 1 allele for each heterozygous individual; the same logic applies for recessive alleles
  • 23. • By convention, if there are 2 alleles at a locus, p and q are used to represent their frequencies • The frequency of all alleles in a population will add up to 1 – For example, p  q  1
  • 25. • For example, consider a population of wildflowers that is incompletely dominant for color – 320 red flowers (CRCR) – 160 pink flowers (CRCW) – 20 white flowers (CWCW) • Calculate the number of copies of each allele – CR  (320  2)  160  800 – CW  (20  2)  160  200
  • 26. • To calculate the frequency of each allele – p  freq CR  800 / (800  200)  0.8 (80%) – q  1  p  0.2 (20%) • The sum of alleles is always 1 – 0.8  0.2  1
  • 27. The Hardy-Weinberg Principle • The Hardy-Weinberg principle describes a population that is not evolving • If a population does not meet the criteria of the Hardy-Weinberg principle, it can be concluded that the population is evolving
  • 28. • The Hardy-Weinberg principle states that frequencies of alleles and genotypes in a population remain constant from generation to generation • In a given population where gametes contribute to the next generation randomly, allele frequencies will not change • Mendelian inheritance preserves genetic variation in a population Hardy-Weinberg Equilibrium
  • 29. • Hardy-Weinberg equilibrium describes the constant frequency of alleles in such a gene pool • Consider, for example, the same population of 500 wildflowers and 1,000 alleles where – p  freq CR  0.8 – q  freq CW  0.2
  • 30. Figure 21.7 Frequencies of alleles Gametes produced p  frequency of CR allele q  frequency of CW allele Alleles in the population Each egg: Each sperm:  0.8  0.2 80% chance 80% chance 20% chance 20% chance
  • 31. • The frequency of genotypes can be calculated – CRCR  p2  (0.8)2  0.64 – CRCW  2pq  2(0.8)(0.2)  0.32 – CWCW  q2  (0.2)2  0.04 • The frequency of genotypes can be confirmed using a Punnett square
  • 32. Figure 21.8 Sperm Eggs 80% CR (p  0.8) 20% CW (q  0.2) p  0.8 q  0.2CR CR CW CW p  0.8 q  0.2 0.64 (p2 ) CR CR 0.16 (pq) CR CW 0.16 (qp) CR CW 0.04 (q2 ) CW CW Gametes of this generation: 64% CR CR , 32% CR CW , and 4% CW CW 64% CR (from CR CR plants) 16% CR (from CR CW plants) 4% CW (from CW CW plants) 16% CW (from CR CW plants) 80% CR  0.8  p 20% CW  0.2  q     64% CR CR , 32% CR CW , and 4% CW CW plants With random mating, these gametes will result in the same mix of genotypes in the next generation:
  • 33. Figure 21.8a Sperm Eggs 80% CR (p  0.8) 20% CW (q  0.2) p  0.8 q  0.2CR CR CW CW p  0.8 q  0.2 0.64 (p2 ) CR CR 0.16 (pq) CR CW 0.16 (qp) CR CW 0.04 (q2 ) CW CW
  • 34. Figure 21.8b Gametes of this generation: 64% CR CR , 32% CR CW , and 4% CW CW 64% CR (from CR CR plants) 16% CR (from CR CW plants) 4% CW (from CW CW plants) 16% CW (from CR CW plants) 80% CR  0.8  p 20% CW  0.2  q     64% CR CR , 32% CR CW , and 4% CW CW plants With random mating, these gametes will result in the same mix of genotypes in the next generation:
  • 35. • If p and q represent the relative frequencies of the only two possible alleles in a population at a particular locus, then – p2  2pq  q2  1 where p2 and q2 represent the frequencies of the homozygous genotypes and 2pq represents the frequency of the heterozygous genotype
  • 37. Conditions for Hardy-Weinberg Equilibrium • The Hardy-Weinberg theorem describes a hypothetical population that is not evolving • In real populations, allele and genotype frequencies do change over time
  • 38. • The five conditions for nonevolving populations are rarely met in nature 1. No mutations 2. Random mating 3. No natural selection 4. Extremely large population size 5. No gene flow
  • 39. • Natural populations can evolve at some loci while being in Hardy-Weinberg equilibrium at other loci • Some populations evolve slowly enough that evolution cannot be detected
  • 40. Applying the Hardy-Weinberg Principle • We can assume the locus that causes phenylketonuria (PKU) is in Hardy-Weinberg equilibrium given that 1. The PKU gene mutation rate is low 2. Mate selection is random with respect to whether or not an individual is a carrier for the PKU allele
  • 41. 3. Natural selection can only act on rare homozygous individuals who do not follow dietary restrictions 4. The population is large 5. Migration has no effect, as many other populations have similar allele frequencies
  • 42. • The occurrence of PKU is 1 per 10,000 births – q2  0.0001 – q  0.01 • The frequency of normal alleles is – p  1 – q  1 – 0.01  0.99 • The frequency of carriers is – 2pq  2  0.99  0.01  0.0198 – or approximately 2% of the U.S. population
  • 43. • Three major factors alter allele frequencies and bring about most evolutionary change – Natural selection – Genetic drift – Gene flow Concept 21.3: Natural selection, genetic drift, and gene flow can alter allele frequencies in a population
  • 44. Natural Selection • Differential success in reproduction results in certain alleles being passed to the next generation in greater proportions • For example, an allele that confers resistance to DDT increased in frequency after DDT was used widely in agriculture
  • 45. Genetic Drift • The smaller a sample, the more likely it is that chance alone will cause deviation from a predicted result • Genetic drift describes how allele frequencies fluctuate unpredictably from one generation to the next • Genetic drift tends to reduce genetic variation through losses of alleles, especially in small populations
  • 46. Figure 21.9-1 CW CW CR CR CR CW CR CR CR CR CR CR CR CR CR CW CR CW CR CW p (frequency of CR )  0.7 q (frequency of CW )  0.3 Generation 1
  • 48. Figure 21.9-3 CW CW CR CR CR CW CR CR CR CR CR CR CR CR CR CW CR CW CR CW CW CW CR CR CR CW CR CR CR CR CR CR CR CR CR CW CR CW CR CW CW CW CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CR CW CW 5 plants leave offspring 2 plants leave offspring p (frequency of CR )  0.7 q (frequency of CW )  0.3 p  0.5 q  0.5 p  1.0 q  0.0 Generation 2 Generation 3Generation 1
  • 49. The Founder Effect • The founder effect occurs when a few individuals become isolated from a larger population • Allele frequencies in the small founder population can be different from those in the larger parent population due to chance
  • 50. The Bottleneck Effect • The bottleneck effect can result from a drastic reduction in population size due to a sudden environmental change • By chance, the resulting gene pool may no longer be reflective of the original population’s gene pool • If the population remains small, it may be further affected by genetic drift
  • 51. Figure 21.10 Original population Surviving population Bottlenecking event (a) By chance, blue marbles are overrepresented in the surviving population. (b) Florida panther (Puma concolor coryi)
  • 52. Figure 21.10a-1 Original population (a) By chance, blue marbles are overrepresented in the surviving population.
  • 53. Figure 21.10a-2 Original population Bottlenecking event (a) By chance, blue marbles are overrepresented in the surviving population.
  • 54. Figure 21.10a-3 Original population Surviving population Bottlenecking event (a) By chance, blue marbles are overrepresented in the surviving population.
  • 55. Figure 21.10b (b) Florida panther (Puma concolor coryi)
  • 56. • Understanding the bottleneck effect can increase understanding of how human activity affects other species
  • 57. Case Study: Impact of Genetic Drift on the Greater Prairie Chicken • Loss of prairie habitat caused a severe reduction in the population of greater prairie chickens in Illinois • The surviving birds had low levels of genetic variation, and only 50% of their eggs hatched
  • 58. Figure 21.11 Pre-bottleneck (Illinois, 1820) Post-bottleneck (Illinois, 1993) Range of greater prairie chicken Illinois 1930–1960s 1993 Greater prairie chicken Kansas, 1998 (no bottleneck) Nebraska, 1998 (no bottleneck) 1,000–25,000 50 75,000– 200,000 5.2 3.7 Location Population size 750,000 Number of alleles per locus Percentage of eggs hatched 93 50 5.8 5.8 99 96 (a) (b)
  • 59. Figure 21.11a Pre-bottleneck (Illinois, 1820) Post-bottleneck (Illinois, 1993) Range of greater prairie chicken Greater prairie chicken (a)
  • 60. • Researchers used DNA from museum specimens to compare genetic variation in the population before and after the bottleneck • The results showed a loss of alleles at several loci • Researchers introduced greater prairie chickens from populations in other states and were successful in introducing new alleles and increasing the egg hatch rate to 90%
  • 61. Effects of Genetic Drift: A Summary 1. Genetic drift is significant in small populations 2. Genetic drift can cause allele frequencies to change at random 3. Genetic drift can lead to a loss of genetic variation within populations 4. Genetic drift can cause harmful alleles to become fixed
  • 62. Gene Flow • Gene flow consists of the movement of alleles among populations • Alleles can be transferred through the movement of fertile individuals or gametes (for example, pollen) • Gene flow tends to reduce genetic variation among populations over time
  • 63. • Gene flow can decrease the fitness of a population • Consider, for example, the great tit (Parus major) on the Dutch island of Vlieland – Immigration of birds from the mainland introduces alleles that decrease fitness in island populations – Natural selection reduces the frequency of these alleles in the eastern population where immigration from the mainland is low – In the central population, high immigration from the mainland overwhelms the effects of selection
  • 64. Figure 21.12 Survivalrate(%) Central population Vlieland, the Netherlands Eastern population NORTH SEA 2 km Population in which the surviving females eventually bred Females born in central population Parus major Central Eastern Females born in eastern population 60 50 40 30 20 10 0
  • 65. • Gene flow can increase the fitness of a population • Consider, for example, the spread of alleles for resistance to insecticides – Insecticides have been used to target mosquitoes that carry West Nile virus and other diseases – Alleles have evolved in some populations that confer insecticide resistance to these mosquitoes – The flow of insecticide resistance alleles into a population can cause an increase in fitness
  • 66. • Gene flow is an important agent of evolutionary change in modern human populations
  • 67. • Evolution by natural selection involves both chance and “sorting” – New genetic variations arise by chance – Beneficial alleles are “sorted” and favored by natural selection • Only natural selection consistently results in adaptive evolution, an increase in the frequency of alleles that improve fitness Concept 21.4: Natural selection is the only mechanism that consistently causes adaptive evolution
  • 68. Natural Selection: A Closer Look • Natural selection brings about adaptive evolution by acting on an organism’s phenotype
  • 69. Relative Fitness • The phrases “struggle for existence” and “survival of the fittest” are misleading as they imply direct competition among individuals • Reproductive success is generally more subtle and depends on many factors
  • 70. • Relative fitness is the contribution an individual makes to the gene pool of the next generation, relative to the contributions of other individuals • Selection indirectly favors certain genotypes by acting directly on phenotypes
  • 71. Directional, Disruptive, and Stabilizing Selection • There are three modes of natural selection – Directional selection favors individuals at one end of the phenotypic range – Disruptive selection favors individuals at both extremes of the phenotypic range – Stabilizing selection favors intermediate variants and acts against extreme phenotypes
  • 72. Figure 21.13 Original population Evolved population Original population Frequencyof individuals Phenotypes (fur color) (a) Directional selection (b) Disruptive selection (c) Stabilizing selection
  • 73. The Key Role of Natural Selection in Adaptive Evolution • Striking adaptations have arisen by natural selection – For example, certain octopuses can change color rapidly for camouflage – For example, the jaws of snakes allow them to swallow prey larger than their heads
  • 74. Figure 21.14 Bones shown in green are movable. Ligament
  • 75. • Natural selection increases the frequencies of alleles that enhance survival and reproduction • Adaptive evolution occurs as the match between an organism and its environment increases • Because the environment can change, adaptive evolution is a continuous, dynamic process
  • 76. • Genetic drift and gene flow do not consistently lead to adaptive evolution, as they can increase or decrease the match between an organism and its environment
  • 77. Sexual Selection • Sexual selection is natural selection for mating success • It can result in sexual dimorphism, marked differences between the sexes in secondary sexual characteristics
  • 79. • Intrasexual selection is competition among individuals of one sex (often males) for mates of the opposite sex • Intersexual selection, often called mate choice, occurs when individuals of one sex (usually females) are choosy in selecting their mates • Male showiness due to mate choice can increase a male’s chances of attracting a female, while decreasing his chances of survival
  • 80. • How do female preferences evolve? • The “good genes” hypothesis suggests that if a trait is related to male genetic quality or health, both the male trait and female preference for that trait should increase in frequency
  • 81. Figure 21.16 SC male gray tree frog LC male gray tree frog Female gray tree frog Recording of SC male’s call Recording of LC male’s call Offspring of SC father SC sperm  Eggs  LC sperm Offspring of LC father Survival and growth of these half-sibling offspring compared Experiment Results
  • 82. Figure 21.16a SC male gray tree frog LC male gray tree frog Female gray tree frog Recording of SC male’s call Recording of LC male’s call Offspring of SC father SC sperm  Eggs  LC sperm Offspring of LC father Survival and growth of these half-sibling offspring compared Experiment
  • 83. The Preservation of Genetic Variation • Neutral variation is genetic variation that does not confer a selective advantage or disadvantage • Various mechanisms help to preserve genetic variation in a population
  • 84. Diploidy • Diploidy maintains genetic variation in the form of hidden recessive alleles • Heterozygotes can carry recessive alleles that are hidden from the effects of selection
  • 85. Balancing Selection • Balancing selection occurs when natural selection maintains stable frequencies of two or more phenotypic forms in a population • Balancing selection includes – Heterozygote advantage – Frequency-dependent selection
  • 86. • Heterozygote advantage occurs when heterozygotes have a higher fitness than do both homozygotes • Natural selection will tend to maintain two or more alleles at that locus • For example, the sickle-cell allele causes deleterious mutations in hemoglobin but also confers malaria resistance
  • 87. Figure 21.17 Distribution of malaria caused by Plasmodium falciparum (a parasitic unicellular eukaryote) Key Frequencies of the sickle-cell allele 10.0–12.5% 12.5% 7.5–10.0% 5.0–7.5% 2.5–5.0% 0–2.5%
  • 88. • Frequency-dependent selection occurs when the fitness of a phenotype declines if it becomes too common in the population • Selection can favor whichever phenotype is less common in a population • For example, frequency-dependent selection selects for approximately equal numbers of “right-mouthed” and “left-mouthed” scale- eating fish
  • 89. Figure 21.18 “Left-mouthed” P. microlepis “Right-mouthed” P. microlepis Sample year Frequencyof “left-mouthed”individuals 1981 ’83 ’85 ’87 ’89 0.5 0 1.0
  • 90. KEY CONCEPT A population shares a common gene pool.
  • 91. Genetic variation in a population increases the chance that some individuals will survive. • Genetic variation leads to phenotypic variation. • Phenotypic variation is necessary for natural selection. • Genetic variation is stored in a population’s gene pool. – made up of all alleles in a population – allele combinations form when organisms have offspring
  • 92. • Allele frequencies measure genetic variation. – measures how common allele is in population – can be calculated for each allele in gene pool
  • 93. Genetic variation comes from several sources. • Mutation is a random change in the DNA of a gene. • Recombination forms new combinations of alleles. – can form new allele – can be passed on to offspring if in reproductive cells – usually occurs during meiosis – parents’ alleles arranged in new ways in gametes
  • 94. Genetic variation comes from several sources. • Hybridization is the crossing of two different species. – occurs when individuals can’t find mate of own species – topic of current scientific research
  • 95. KEY CONCEPT Populations, not individuals, evolve.
  • 96. Natural selection acts on distributions of traits. • A normal distribution graphs as a bell-shaped curve. • Traits not undergoing natural selection have a normal distribution. – highest frequency near mean value – frequencies decrease toward each extreme value
  • 97. Natural selection can change the distribution of a trait in one of three ways. • Microevolution is evolution within a population. – observable change in the allele frequencies – can result from natural selection
  • 98. • Natural selection can take one of three paths. – Directional selection favors phenotypes at one extreme.
  • 99. – Stabilizing selection favors the intermediate phenotype. • Natural selection can take one of three paths.
  • 100. • Natural selection can take one of three paths. – Disruptive selection favors both extreme phenotypes.
  • 101. KEY CONCEPT Natural selection is not the only mechanism through which populations evolve.
  • 102. Gene flow is the movement of alleles between populations. • Gene flow occurs when individuals join new populations and reproduce. • Gene flow keeps neighboring populations similar. • Low gene flow increases the chance that two populations will evolve into different species. bald eagle migration
  • 103. Genetic drift is a change in allele frequencies due to chance. • Genetic drift causes a loss of genetic diversity. • It is most common in small populations. • A population bottleneck can lead to genetic drift. – It occurs when an event drastically reduces population size. – The bottleneck effect is genetic drift that occurs after a bottleneck event.
  • 104. • The founding of a small population can lead to genetic drift. – It occurs when a few individuals start a new population. – The founder effect is genetic drift that occurs after start of new population.
  • 105. • Genetic drift has negative effects on a population. – less likely to have some individuals that can adapt – harmful alleles can become more common due to chance
  • 106. Sexual selection occurs when certain traits increase mating success. • Sexual selection occurs due to higher cost of reproduction for females. – males produce many sperm continuously – females are more limited in potential offspring each cycle
  • 107. • There are two types of sexual selection. – intrasexual selection: competition among males – intersexual selection: males display certain traits to females
  • 108. KEY CONCEPT New species can arise when populations are isolated.
  • 109. The isolation of populations can lead to speciation. • Populations become isolated when there is no gene flow. – Isolated populations adapt to their own environments. – Genetic differences can add up over generations.
  • 110. • Reproductive isolation can occur between isolated populations. – members of different populations cannot mate successfully – final step to becoming separate species • Speciation is the rise of two or more species from one existing species.
  • 111. Populations can become isolated in several ways. • Behavioral barriers can cause isolation. – called behavioral isolation – includes differences in courtship or mating behaviors
  • 112. • Geographic barriers can cause isolation. – called geographic isolation – physical barriers divide population • Temporal barriers can cause isolation. – called temporal isolation – timing of reproductive periods prevents mating
  • 114. Evolution through natural selection is not random. • Natural selection can have direction. • The effects of natural selection add up over time.
  • 115. • Convergent evolution describes evolution toward similar traits in unrelated species.
  • 116. • Divergent evolution describes evolution toward different traits in closely related species. How do convergent and divergent evolution illustrate the directional nature of natural selection? ancestor red foxkit fox
  • 117. Species can shape each other over time. • Two or more species can evolve together through coevolution. – evolutionary paths become connected – species evolve in response to changes in each other
  • 118. • Coevolution can occur in beneficial relationships.
  • 119. • Coevolution can occur in competitive relationships, sometimes called evolutionary.
  • 120. Species can become extinct. • Extinction is the elimination of a species from Earth. • Background extinctions occur continuously at a very low rate. – occur at roughly the same rate as speciation – usually affects a few species in a small area – caused by local changes in environment
  • 121. • Background extinctions occur continuously at a very low rate. – occur at roughly the same rate as speciation – usually affects a few species in a small area – caused by local changes in environment
  • 122. • Mass extinctions are rare but much more intense. – destroy many species at global level – thought to be caused by catastrophic events – at least five mass extinctions in last 600 million years
  • 123. Speciation often occurs in patterns. • A pattern of punctuated equilibrium exists in the fossil record. – theory proposed by Eldredge and Gould in 1972 – episodes of speciation occur suddenly in geologic time – followed by long periods of little evolutionary change – revised Darwin’s idea that species arose through gradual transformations
  • 124. • Many species evolve from one species during adaptive radiation. – ancestral species diversifies into many descendent species – descendent species usually adapted to wide range of environments
  • 125. Moral of this story : Survival of the fittest…….
  • 126. References James F. Hancock 2012 Plant Evolution and the Origin of Crop Species 3rd eds CABI Publishing John D. Thompson 2005 Plant Evolution in the Mediterranean Oxford University Press Miller AJ. 2007. Crop Plants: Evolution. John Wiley & Sons. Peter R. Bell, Alan R. Hemsley 1992 Green Plants Their Origin and Diversity Cambridge University Press