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Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

PowerPoint Lectures forBiology, Seventh Edition

 Neil Campbell and Jane Reece

Lectures by Chris Romero

Chapter 23

he E!olution o" Populations

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• Overview: The Smallest Unit of Evolution

• One common misconception about evolution is

that individual organisms evolve, in the

arwinian sense, during their lifetimes

• !atural selection acts on individuals, but

populations evolve

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• "enetic variations in populations

 #  #ontribute to evolution

Figure 23.1

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• #oncept $%&': Population genetics provides a

foundation for stud(ing evolution• )icroevolution

 #  *s change in the genetic ma+eup of a

population from generation to generation

Figure 23.2

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The Modern Synthesis

• Population genetics

 #  *s the stud( of how populations changegeneticall( over time

 #  econciled arwin-s and )endel-s ideas

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• The modern s(nthesis

 #  *ntegrates )endelian genetics with thearwinian theor( of evolution b( natural

selection

 #  .ocuses on populations as units of evolution

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Gene Pools and Allele Frequencies

•  / population

 #  *s a locali0ed group of individuals that are capable ofinterbreeding and producing fertile offspring

)/P

 /E/

      /      L      /      S      1      /

      #       /      !      /            /

2eaufor t Sea

Porcupine

herd r ange

• 

.airban+s

• 

3hitehorse

.ort(mile

herd range

!   O  ,  T   4   3   

E   S  T   

T   E   ,  ,  *  T   O  ,  *  E   S  

     /     L     /     S     1     /

     5     U     1     O     !

Figure 23.3

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• The gene pool

 #  *s the total aggregate of genes in a populationat an( one time

 #  #onsists of all gene loci in all individuals of the

population

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The Hardy-Weinber Theorem

• The 4ard(63einberg theorem

 #  escribes a population that is not evolving

 #  States that the fre7uencies of alleles and

genot(pes in a population-s gene pool remain

constant from generation to generationprovided that onl( )endelian segregation and

recombination of alleles are at wor+

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• )endelian inheritance

 #  Preserves genetic variation in a population

Figure 23.4

"eneration

'

C R C R 

genot(pe

C W C W 

genot(pePlants mate

 /ll C R C W 

8all pin+ flowers9

;< C R 

gametes;< C W 

gametes

#ome together at random

"eneration

$

"eneration

%

"eneration

=

$< C R C R  ;< C R C W  $< C W C W 

;< C R 

gametes;< C W 

gametes

#ome together at random

$< C R C R  ;< C R C W  $< C W C W 

 /lleles segregate, and subse7uent

generations also have three t(pes

of flowers in the same proportions

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 Preservation of Allele Frequencies

• *n a given population where gametes contribute

to the ne>t generation randoml(, allelefre7uencies will not change

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 Hardy-Weinberg quilibrium

• 4ard(63einberg e7uilibrium

 #  escribes a population in which randommating occurs

 #  escribes a population where allele

fre7uencies do not change

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•  / population in 4ard(63einberg e7uilibrium

Figure 23.5

"ametes for each generation are drawn at random from

the gene pool of the previous generation:

?;< C R  8 p @ ;&?9 $;< C W  8q @ ;&$9

SpermC R 

8?;<9C W 

8$;<9

 p2 

A=<C R C R 

'A<C R C W 

'A<C R C W 

=<C W C W 

qp

    C    R

   8   ?   ;   <   9

   E  g  g  s

    C    W

   8   $   ;   <   9

 pq

*f the gametes come together at random, the genot(pe

fre7uencies of this generation are in 4ard(63einberg e7uilibrium:

q$

A=< C R C R , %$< C R C W , and =< C W C W 

"ametes of the ne>t generation:

A=< C R  from

C R C R  homo0(gotes

'A< C R  from

C R C W  homo0(gotesB @ ?;< C R  @ ;&? @ p

'A< C W  from

C R C W  hetero0(gotesB @ $;< C W  @ ;&$ @ q

3ith random mating, these gametes will result in the same

mi> of plants in the ne>t generation:

A=< C R 

C R 

, %$< C R 

C W 

 and =< C W 

C W 

 plants

 p2 

=< C W  from

C W C W  homo0(gotes

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• *f p and q represent the relative fre7uencies of

the onl( two possible alleles in a population ata particular locus, then

 #   p$ B $ pq B q$ @ '

 #   /nd p$ and q2  represent the fre7uencies of thehomo0(gous genot(pes and $ pq represents

the fre7uenc( of the hetero0(gous genot(pe

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Conditions for Hardy-Weinberg quilibrium

• The 4ard(63einberg theorem

 #  escribes a h(pothetical population

• *n real populations

 #   /llele and genot(pe fre7uencies do changeover time

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• The five conditions for non6evolving

populations are rarel( met in nature #  E>tremel( large population si0e

 #  !o gene flow

 #  !o mutations

 #  andom mating

 #  !o natural selection

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 Population !enetics and Human Healt"

• 3e can use the 4ard(63einberg e7uation

 #  To estimate the percentage of the humanpopulation carr(ing the allele for an inherited

disease

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• #oncept $%&$: )utation and se>ual

recombination produce the variation thatma+es evolution possible

• Two processes, mutation and se>ual

recombination #  Produce the variation in gene pools that

contributes to differences among individuals

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Mutation

• )utations

 #   /re changes in the nucleotide se7uence of !/

 #  #ause new genes and alleles to arise

Figure 23.6

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 Point #utations

•  / point mutation

 #  *s a change in one base in a gene

 #  #an have a significant impact on phenot(pe

 #  *s usuall( harmless, but ma( have an adaptiveimpact

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 #utations $"at Alter !ene Number or %equence

• #hromosomal mutations that affect man( loci

 #   /re almost certain to be harmful

 #  )a( be neutral and even beneficial

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• "ene duplication

 #  uplicates chromosome segments

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 #utation Rates

• )utation rates

 #  Tend to be low in animals and plants

 #   /verage about one mutation in ever( ';;,;;;

genes per generation

 #   /re more rapid in microorganisms

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Se!ual Recombination

• *n se>uall( reproducing populations, se>ual

recombination #  *s far more important than mutation in

producing the genetic differences that ma+e

adaptation possible

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• #oncept $%&%: !atural selection, genetic drift,

and gene flow can alter a population-s geneticcomposition

• Three maCor factors alter allele fre7uencies and

bring about most evolutionar( change #  !atural selection

 #  "enetic drift

 #  "ene flow

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"atural Selection

• ifferential success in reproduction

 #  esults in certain alleles being passed to thene>t generation in greater proportions

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Genetic #ri$t

• Statisticall(, the smaller a sample

 #  The greater the chance of deviation from apredicted result

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• "enetic drift

 #  escribes how allele fre7uencies can fluctuateunpredictabl( from one generation to the ne>t

 #  Tends to reduce genetic variation

Figure 23.7

C R C R 

C R C W 

C R C R 

C W C W  C R C R 

C R C W 

C R C W 

C R C W C R C R 

C R C R 

Onl( of 

'; plants

leave

offspring

C W C W  C R C R 

C R C W 

C R C R    C W C W 

C R C W 

C W C W  C R C R 

C R C W  C R C W 

Onl( $ of 

'; plants

leave

offspring

C R C R 

C R C R  C R C R 

C R C R C R C R 

C R C R 

C R C R 

C R C R 

C R C R C R C R 

Generation 2

 p @ ;&

q @ ;&

Generation 3

 p @ '&;

q @ ;&;

Generation 1

 p 8fre7uenc( of C R 9 @ ;&D

q 8fre7uenc( of C W 9 @ ;&%

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$"e &ottlenec' ffect 

• *n the bottlenec+ effect

 #   / sudden change in the environment ma(drasticall( reduce the si0e of a population

 #  The gene pool ma( no longer be reflective of

the original population-s gene pool

Original

population

2ottlenec+ing

eventSurviving

population

Figure 23.8 A

(a) $ha%ing just a "e& marbles through the

narro& nec% o" a bottle is analogous to a

drastic reduction in the si'e o" a population

a"ter some en!ironmental disaster. By chance,

 blue marbles are o!er(represented in the ne&

 population and gold marbles are absent.

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• Understanding the bottlenec+ effect

 #  #an increase understanding of how humanactivit( affects other species

Figure 23.8 B

(b) $imilarly, bottlenec%ing a population

o" organisms tends to reduce genetic

!ariation, as in these northern

elephant seals in Cali"ornia that &ereonce hunted nearly to e)tinction.

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$"e Founder ffect 

• The founder effect

 #  Occurs when a few individuals becomeisolated from a larger population

 #  #an affect allele fre7uencies in a population

G Fl

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Gene Flo%

• "ene flow

 #  #auses a population to gain or lose alleles

 #  esults from the movement of fertile

individuals or gametes

 #  Tends to reduce differences between

populations over time

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• #oncept $%&=: !atural selection is the primar(

mechanism of adaptive evolution

• !atural selection

 #   /ccumulates and maintains favorable

genot(pes in a population

G ti & i ti

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Genetic &ariation

• "enetic variation

 #  Occurs in individuals in populations of allspecies

 #  *s not alwa(s heritable

Figure 23.9 A, B

8a9 )ap butterflies that

emerge in spring:

orange and brown

8b9 )ap butterflies that

emerge in late summer:

blac+ and white

( i i Wi "i P l i

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(ariation Wit"in a Population

• 2oth discrete and 7uantitative characters

 #  #ontribute to variation within a population

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• iscrete characters

 #  #an be classified on an either6or basis

• uantitative characters

 #  Far( along a continuum within a population

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• Pol(morphism

• Phenot(pic pol(morphism

 #  escribes a population in which two or more

distinct morphs for a character are each

represented in high enough fre7uencies to bereadil( noticeable

• "enetic pol(morphisms

 #   /re the heritable components of characters

that occur along a continuum in a population

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• )easuring "enetic Fariation

• Population geneticists

 #  )easure the number of pol(morphisms in a

population b( determining the amount of

hetero0(gosit( at the gene level and themolecular level

•  /verage hetero0(gosit(

 #  )easures the average percent of loci that are

hetero0(gous in a population

( i ti & t P l ti

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(ariation &et)een Populations

• )ost species e>hibit geographic variation

 #  ifferences between gene pools of separatepopulations or population subgroups

' $&= %&'= &'? A D&'

GG'H'%&'D';&'AH&'$?&''

' $&'H %&? =&'A &'= A&D

GG'&'?'%&'D''&'$H&';Figure 23.1

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• Some e>amples of geographic variation occur

as a cline, which is a graded change in a trait

along a geographic a>is

Figure 23.11

!"#!$%&!'  esearchers observed that the average si0e

of (arrow plants 8 Achillea9 growing on the slopes of the Sierra

!evada mountains graduall( decreases with increasing

elevation& To eliminate the effect of environmental differences

at different elevations, researchers collected seeds

from various altitudes and planted them in a common

garden& The( then measured the heights of theresulting plants&

$!*+  The average plant si0es in the common

garden were inversel( correlated with the altitudes at

which the seeds were collected, although the height

differences were less than in the plants- natural

environments&

C'C+*%'  The lesser but still measurable clinal variation

in (arrow plants grown at a common elevation demonstrates the

role of genetic as well as environmental differences&

   )  e  a  n   h  e   i  g   h   t   8  c  m   9

   /   t   i   t  u   d  e   8  m   9

4eights of (arrow plants grown in common garden

Seed collection sites

Sierra !evada

ange"reat 2asin

Plateau

A Closer Loo' at "atural Selection

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A Closer Loo' at "atural Selection

• .rom the range of variations available in a

population

 #  !atural selection increases the fre7uencies of

certain genot(pes, fitting organisms to their

environment over generations

volutionary Fitness

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 volutionary Fitness

• The phrases Istruggle for e>istenceJ and

Isurvival of the fittestJ

 #   /re commonl( used to describe natural

selection

 #  #an be misleading

• eproductive success

 #  *s generall( more subtle and depends on man(factors

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• .itness

 #  *s the contribution an individual ma+es to thegene pool of the ne>t generation, relative to

the contributions of other individuals

• elative fitness

 #  *s the contribution of a genot(pe to the ne>t

generation as compared to the contributions of

alternative genot(pes for the same locus

*irectional *isruptive and %tabili,ing %election

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 *irectional+ *isruptive+ and %tabili,ing %election

• Selection

 #  .avors certain genot(pes b( acting on thephenot(pes of certain organisms

• Three modes of selection are

 #  irectional

 #  isruptive

 #  Stabili0ing

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• irectional selection

 #  .avors individuals at one end of thephenot(pic range

• isruptive selection

 #  .avors individuals at both e>tremes of the

phenot(pic range

• Stabili0ing selection

 #  .avors intermediate variants and acts against

e>treme phenot(pes

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• The three modes of selection

Fig 23.12 A-C

(a) ire/tiona0 e0e/tion shifts the overall

ma+eup of the population b( favoring

variants at one e>treme of the

distribution& *n this case, dar+er mice are

favored because the( live among dar+

roc+s and a dar+er fur color conceals them

from predators&

(b) iruptie e0e/tion favors variants

at both ends of the distribution& These

mice have coloni0ed a patch( habitat

made up of light and dar+ roc+s, with the

result that mice of an intermediate color are

at a disadvantage&

(/) tabi0iing e0e/tion removes

e>treme variants from the population

and preserves intermediate t(pes& *f 

the environment consists of roc+s of 

an intermediate color, both light and

dar+ mice will be selected against&

Phenot(pes 8fur color9

Original population

Original

population

Evolved

population

   .  r  e  7  u  e  n  c  (

  o   f   i  n   d   i  v   i   d  u  a   l  s

The Preser(ation o$ Genetic &ariation

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The Preser(ation o$ Genetic &ariation

• Farious mechanisms help to preserve genetic

variation in a population

#i)loidy

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#i)loidy

• iploid(

 #  )aintains genetic variation in the form ofhidden recessive alleles

*alancin Selection

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*alancin Selection

• 2alancing selection

 #  Occurs when natural selection maintainsstable fre7uencies of two or more phenot(pic

forms in a population

 #  Leads to a state called balanced pol(morphism

Hetero+yote Ad(antae

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Hetero+yote Ad(antae

• Some individuals who are hetero0(gous at a

particular locus

 #  4ave greater fitness than homo0(gotes

• !atural selection

 #  3ill tend to maintain two or more alleles at that

locus

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• The sic+le6cell allele

 #  #auses mutations in hemoglobin but alsoconfers malaria resistance

 #  E>emplifies the hetero0(gote advantage

Figure 23.13

.re7uencies of thesic+le6cell allele

;K$&<

$&K&;<

&;KD&<

D&K';&;<

';&;K'$&<

'$&<

istribution of 

malaria caused b(

Plasmodium falciparum

8a proto0oan9

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• .re7uenc(6ependent Selection

• *n fre7uenc(6dependent selection

 #  The fitness of an( morph declines if it becomes

too common in the population

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•  /n e>ample of fre7uenc(6dependent selection

   P   h  e  n  o   t  (  p   i  c

   d   i  v  e  r  s   i   t  (

Figure 23.14

#arenta0 popu0ation ap0e

!perienta0 group ap0e

Plain bac+ground Patterned bac+ground

On pec+ing a moth image

the blue Ca( receives a

food reward& *f the bird does not detect a moth

on either screen, it pec+s

the green circle to continue

to a new set of images 8a

new feeding opportunit(9&

;&;A

;&;

;&;=

;&;%

;&;$

; $; =; A; ?; ';;"eneration number 

.re7uenc(6

independent control

Neutral (ariation

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 Neutral (ariation

• !eutral variation

 #  *s genetic variation that appears to confer noselective advantage

Se!ual Selection

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Se!ual Selection

• Se>ual selection

 #  *s natural selection for mating success

 #  #an result in se>ual dimorphism, mar+ed

differences between the se>es in secondar(

se>ual characteristics

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• *ntrase>ual selection

 #  *s a direct competition among individuals ofone se> for mates of the opposite se>

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• *nterse>ual selection

 #  Occurs when individuals of one se> 8usuall(females9 are choos( in selecting their mates

from individuals of the other se>

 #  )a( depend on the showiness of the male-sappearance

Figure 23.15

The ,(olutionary ,nima o$ Se!ual Re)roduction

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The ,(olutionary ,nima o$ Se!ual Re)roduction

• Se>ual reproduction

 #  Produces fewer reproductive offspring than ase>ualreproduction, a so6called reproductive handicap

Figure 23.16

Aeua0 reprou/tion

.emale

eua0 reprou/tion

.emale

)ale

"eneration '

"eneration $

"eneration %

"eneration =

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• *f se>ual reproduction is a handicap, wh( has it

persistedM

 #  *t produces genetic variation that ma( aid in

disease resistance

Why "atural Selection Cannot Fashion Per$ect ranisms

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y

• Evolution is limited b( historical constraints

•  /daptations are often compromises

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• #hance and natural selection interact

• Selection can onl( edit e>isting variations