Chapter 13 PHENOTYPIC EVOLUTION

Summary of Chapter 13: EVOLUTION, D. Futuyma, 2

n
E., 2!!"
Variation in most phenotypic characters is continuous or quantitative, and it is based on effects of several
or many variable gene loci, as well as those of the environment.
Height, finger length, eye color, skin color, etc.
uantitative genetics studies continuous characters and the mechanisms that produce them.
!t includes traits that are determined by several genes" polygenic traits.
#hese traits may have a strong environmental component.
#$uant%tat%&e 'enet%() ea*) +%th the 'enet%() of (ont%nuou)*y &ary%n' (hara(ter). ,ather than (on)%er%n' (han'e) %n the
fre-uen(%e) of )pe(%f%( a**e*e) of 'enotype), -uant%tat%&e 'enet%() )ee.) to /-uant%fy/ (han'e) %n the fre-uen(y %)tr%0ut%on of
tra%t) that (annot ea)%*y 0e p*a(e %n %)(rete phenotyp%( (*a))e). The rea)on for the (ont%nuou) &ar%at%on %) u)ua**y that the tra%t)
are polygenic 1(ontro**e 0y many 'ene)2 an there are environmental effe(t) that a*ter the phenotyp%( )tate of ea(h %n%&%ua*.3
http"$$biomed.brown.edu$Courses$%!&'($1).uan.genetics.H#*+
GENETIC ARCHITECTURE OF PHENOTYPIC TRAITS
Genetic architecture refers to the genetic basis of a trait and its relationship to other traits.
,enetic architecture has several components"
-umber of loci that contribute to its development, variation within species and among species or
populations.
*agnitude of the effect on the trait of different alleles at each locus . the allelic effect.
#he additive effect and dominance at the various loci.
#he synergistic effect among loci . epistasis.
#he pleitrpic effect of the loci . the e/tent to which individual genes affect more than one
trait.
#he field of !uantitati"e #enetics analy0es quantitative characters. !t studies the evolution of
morphology, life history characteristics, behavior, etc.
uantitative, multifactorial or polygenic inheritance refers to phenotypic characteristics that are the result
of two or more genes and their interaction with the environment.
1olygenic traits do not follow *endelian inheritance, and the phenotypes vary along a gradient depicted
by a bell shape curve.
2 !uantitati"e trait lcus 3$TL4 is a region of 5-2 that is associated with a particular phenotypic trait
6 these #+s are often found on different chromosomes.
2 #+ is not necessarily a single gene, but rather a chromosome interval that may include many genes of
which one or more affect the character under study.
#+s are loci that influence a quantitative trait.
2 !uantitati"e trait lcus 3$TL4 is a region of 5-2 that is associated with a particular phenotypic trait
6 these #+s are often found on different chromosomes.
7nowing the number of #+s that e/plains variation in the phenotypic trait tells us about the genetic
architecture of a trait.
!t may tell us that plant height is controlled by many genes of small effect, or by a few genes of
large effect.
8ome #+s have large effects while other have small effects.
*any #+s have both additive effects on the phenotype and strong epistatic interaction.
8trong epistatic effect means that the 9oint effect of some pairs of loci differed from the sum of their
individual effects.
8ome #+s show additive effect on the phenotype.
1henotypic effects can differ among the se/es other effects depend on the temperature at which the
organisms are raised
$TL %appin# is the statistical study of the alleles that occur in a locus and the phenotypes 3physical
forms or traits4 that they produce.
#+ mapping is the collective name for a suite of related techniques that employ marker loci to
scan chromosomes and identify regions containing genes that contribute to a quantitative trait.
3:reeman and Herron, ;))<4
2 marker is segment of 5-2 that is significantly more likely to co6occur with the trait than e/pected by
chance that is a marker that has a statistical association with the trait.
Can&i&ate #enes are genes thought to be involved in the evolution of a phenotypic trait based on its
mutant phenotype of the protein they encode.
CO'PONENTS OF PHENOTYPIC VARIATION
Variance" average of the squared deviations of observations from the arithmetic mean of a sample 3V4.
Stan&ar& &e"iatin" the square root of the variance, s = >V.
http"$$www.chssc.salford.ac.uk$health8ci$resmeth;)))$resmeth$variance.htm
!f a variable has a normal 3bell shaped4 frequency distribution, about ?(@ of the observations lie within
one standard deviation on either side of the mean, A?@ within two standard deviations, and AA.<@ within
three.
Variation 3variance4 in a phenotypic trait 3V14 may include genetic variance 3V,4 and variance due to
environmental factors 3VB4.
#he phenotype variation among individuals with the same genotype is the environmental variance, VB.
,enetic variance may include both a&&iti"e #enetic "ariance 3V24 due to the additive effects of alleles
and non6additive genetic variance due to dominance and epistasis.
2dditive effects of alleles are responsible for the degree of similarity between parents and offspring and
are the basis for response to selection within population.
&nly the additive variance creates a correlation between parents and offspring.
&nly V2 is affected by selection.
Phent(pic "ariance 3V14 is variance of the phenotype 3physical and biochemical characteristics4 that
results from the sum of the #ent(pic 3V,4 and en"irn%ental 3VB4 variances.
V1 = V, C VB
A&&iti"e #enetic "ariance" Dhen the hetero0ygote lies at midpoint between two homo0ygous,
inheritance is the additive.
2121 212; 2;2;
61 ) C1
#he additive genetic variance depends both on the magnitude of the a&&iti"e effects f alleles on the
phenotype and on the #ent(pe fre!uencies.
!f a genotype is too frequent, most individuals are close to the average phenotype and the
variance is lower.
!f several genotypes have similar frequencies, the variance will be greater.
Dhen several loci contribute additively to the phenotype, the average phenotype of any particular
genotype is the sum of the phenotypic values of each of the loci.
#he additive genetic variance plays a key role in evolutionary theory because the additive effects of
alleles are responsible for the degree of similarity between parents and offspring and, therefore, are the
basis for response to selection within populations.
Respnse t selectin is the change in the average genotype after selection in the previous generation.
8election among phenotypically different parents is reflected in the mean phenotype of the ne/t
generation.
#he proportion of the genetic variance to the total variance is herita)ilit(.
Heritability does not indicate the degree to which a trait is geneticE it measures the proportion of the
phenotypic variance that is the result of genetic factors.
#he heritability is determined by the additive genetic variance 3V24, which depends on allele frequenciesE
and by the environmental variance 3VB4, which depends in part on how variable the environmental factors
are that affect the development or e/pression of the character.
#wo specific types of heritability can be estimated.
#he broad6sense heritability is the ratio of total #enetic "ariance to total phent(pic "ariance.
H
*
+ VG,VP
#he narrow6sense heritability is the ratio of a&&iti"e #enetic "ariance to the total phent(pic
"ariance.
h
*
N + VA,VP where V1 = V,CVB and V, is the sum of V2 and non6additive
genetic components.
#he estimate heritability is valid only for the population in which it was measured, and only in the
environment in which it was measured. #his is because frequencies vary between populations and so do
environmental conditions.
Narr- sense herita)ilit( -uant%f%e) on*y the port%on of the phenotyp%( &ar%at%on that %) a%t%&e 1a**e*%(2
0y nature4 5hen %ntere)te %n %mpro&%n' *%&e)to(. &%a art%f%(%a* )e*e(t%on, for e6amp*e, .no+%n' the
narro+7)en)e her%ta0%*%ty of the tra%t of %ntere)t +%** a**o+ pre%(t%n' ho+ mu(h the mean of the tra%t +%**
%n(rea)e %n the ne6t 'enerat%on a) a fun(t%on of ho+ mu(h the mean of the )e*e(te parent) %ffer) from
the mean of the popu*at%on from +h%(h the )e*e(te parent) +ere (ho)en. The o0)er&e re)pon)e to
)e*e(t%on *ea) to an e)t%mate of the narro+7)en)e her%ta0%*%ty 1(a**e realized heritability2.
http"$$en.wikipedia.org$wiki$Heritability
LIN.AGE /ISE$UILI0RIU'
+inkage disequilibrium is a term used in population genetics.
Dhen alleles at two loci are randomly associated with each other, they are said to in a state of lin1a#e
e!uili)riu%E when they are non6randomly associated, they are in lin1a#e &ise!uili)riu%
Lin1a#e &ise!uili)riu%" #he association of two alleles at two or more loci more or less frequently than
predicted by their individual frequencies
#he loci in linkage disequilibrium are usually located in different chromosomes.
+inkage disequilibrium is generally caused by selection in the form of epistatic fitness interactions
between genes, genetic linkage and the rate of recombination, random drift or non6random mating, and
population structure.
Epistasis occurs when the effect of a gene is affected by the activity of other genes located in different
loci.
2 gene produces a protein that prevents transcription of another gene.
#he coefficient of linkage disequilibrium is the measure of the deviation from the two6locus equilibrium.
1ositive linkage disequilibrium increases the frequency of phenotypic variance.
#he level of linkage disequilibrium declines due to recombination in the double hetero0ygotes with the
passage of generations.
+inkage disequilibrium is caused by"
1. -atural selection. !f selection favors individuals with particular combinations of alleles, then it
produces linkage disequilibrium. !f two or more gene combinations are much fitter than recombinant
genotypes linkage disequilibrium will be favored.
;. -on6random mating.
3. Dhen a new mutation arises, the single copy is necessarily associated with specific alleles at other loci
on the chromosome, and therefore is in linkage disequilibrium with those alleles. #he copies of this
mutation in subsequent generations will retain this association until it is broken down by recombination.
'. #he linkage equilibrium has not yet been reached. !t takes a number of generations for recombination
to do its randomi0ing work and, particularly for tightly linked genesE linkage disequilibrium can persist
for some time.
F. Gecombination is very slow or non6e/istent. Gecombination tends to randomi0e genotypes at one locus
with respect to genotypes at another. !t decreases the frequency of overrepresented haplotypes and
increases the frequencies of underrepresented haplotypes.
?. Gandom drift processes can cause persistent linkage disequilibrium. !f random sampling produces by
chance an e/cess of a haplotype in a generation, linkage disequilibrium will have arisen. 2ny haplotype
could be HfavoredH by chance, so the disequilibrium is equally likely to have 5 I ) or 5 J ).
EVOLUTION OF $UANTITATIVE CHARACTERS
GENETIC VARIANCE IN NATURAL POPULATIONS
Herita)ilit( is determined by the additive genetic variance 3VA2, which depends on allele frequenciesE
and by the environmental variance 3VE4, which depends in part on how variable the environmental factors
are that affect the development or e/pression of the character. Vp is the phenotypic variance.
h
*
+ VA,VP
Dhere V1 = V, C VBE and V, is the sum of V2 and non6additive genetic components.
#he causes of genetic variation in natural populations are uncertain, but input by mutation may balance
losses due to selection and genetic drift.
#he paucity of genetic variation would be a genetic constraint that could affect the direction of evolution
or prevent adaptation altogether.
E3a%ple" 8ome grasses growing near mines have developed tolerance to high copper and 0inc
concentration in the soilE other grass species have not and disappeared from the contaminated
areas. B/periments showed that seeds from plants growing in normal soil produced a small
number of tolerant seedlings in the species that evolved tolerance in contaminated soilsE no
tolerant seedlings were found in those species that had not formed tolerant populations in
contaminated soil. #he genetic variation that allowed for tolerance was apparently missing in the
intolerant species.
GENETIC /RIFT OR NATURAL SELECTION
2 -BK#G2+ *&5B+ &: #HB BV&+K#!&- &: K2-#!#2#!VB CH2G2C#BG8
uantitative characters will evolve by genetic drift if alleles at the underlying loci are selectively neutral.
8election may be inferred if the data do not fit a model of neutral evolution.
-eutral traits may have a pleiotropic effect on other traits and, therefore, affect fitness.
2t equilibrium, mutation is balanced by genetic drift, the genetic variance and heritability should
theoretically reach a stable value, which should be quite high if the effective population si0e is large.
2s mutations are fi/ed by genetic drift, the mean will fluctuate randomly.
*utation and genetic drift can cause divergence between two isolated populations that originated from a
uniform ancestral population.
2llele frequency changes by genetic drift transpire more rapidly in smaller populations and will be
noticeable in relatively short time.
!f alleles that contribute to variation in a polygenic trait are selectively neutral, variation and evolution of
the trait are affected only by mutation 3which increases variation4 and genetic drift 3which erodes it4.
&ver a long time, if a trait has diverged in populations that originated from a common ancestral
population much faster than the e/pected neutral rate, it is likely to have evolved by directional selection.
!f it has changed much more slowly than the e/pected neutral rate, it is likely that stabili0ing selection has
operated either on the trait itself or on other characteristics that are pleitropically affected by the same
genes.
8tudies on the rate of evolution show that many features fluctuate rather rapidly, but show little net
change, so that the rate of evolution over the long term is much less than it is over shorter intervals of
time.
#he rate of evolution over the long term is much less than it is over short periods of time suggesting that
stabili0ing or fluctuating selection has been in operation.
!n stabili0ing selection, the intermediate phenotypes are preferred instead of the most e/tremes and
variation decreases.
NATURAL SELECTION ON $UANTITATIVE TRAITS
45 RESPONSE TO /IRECTIONAL SELECTION
#he response of a quantitative character to selection depends on the heritability of the character and the
selection differential.

#he selectin &ifferential 6S2 is the difference between the mean of a population and the mean of the
individuals selected to be parents of the ne/t generation.
#he respnse t selectin 6R2 is the change in the mean phenotype after selection.
Herita)ilit( describes how a trait will respond to selection.
B/ample" #he trait to be selected is the length of the tail in a population of ratsE the trait has a normal
frequency distribution 7 3bell6shaped curve4. See F%'. 13.8.
2 normal distribution is e/pected if a large number of loci, all with relatively small effects on the
character, freely recombine.
#he e/perimenter imposes a selection for tail length by breeding only those rats with a tail longer
than a certain value.
#he mean tail length of the selected parents differs from the mean of the entire populationE this is
the selection differential, S.
#he average tail length among the offspring of the selected parents differs from that on the
parental generation as a whole by an amount R, the response to selection.
#he magnitude of R is proportionate to the heritability of the trait.
2s selection proceeds, it increases the frequencies of those alleles that produce phenotypes closer to the
optimum value.
2s those frequencies increase, combination of alleles at different loci 3multilocus genotypes4 that were
e/tremely rare, become more common.
!n this way, phenotypes arise that were absent before.
#he mean of a polygenic character shifts beyond the original range of variation as directional selection
proceeds, even if no further mutations occur.
2s selection proceeds, LlowM alleles are removed and additive genetic variation is reduced.
2s the V2 decreases, the heritability decreases" h
*
+ VA,VP
1rolonged directional selection should ultimately fi/ all favored alleles, eliminating genetic
variation.
2lleles with the most favorable effects on the phenotype are likely to be fi/ed first.
Dill selection come to a haltN 1robably not because mutation is constantly introducing a trickle of
new alleles each with different additive effects.
*utational variance, the infusion of new additive genetic variance by mutation, is on the order of
1)
63
/ VB per generation. 2 fully homo0ygous population could attain a heritability of ).F in one
thousand generations.
!f mutations have harmful pleiotropic effects and have a net selective disadvantage, the LusableM
3advantageous4 mutational variance would be much less.
Knder this view a continual mutation6selection balance might e/plain the gradual changes in
phenotype seen over long evolutionary times.
*5 RESPONSES TO ARTIFICIAL SELECTION
2rtificial selection is the intentional selection of trait or combination of traits by humans.
#he criterion for survival and reproduction is the trait chosen, rather than fitness as determined by the
entire genotype.
#his is in contrast with natural selection, which focuses on the organism ability to survive and reproduce
in its natural habitat, the overall fitness.
2n e/periment by Ooo 31A()4 in which he selected for an increased number of bristles in 5rosophila
melanogaster showed that artificial selection has accomplished an enormous evolutionary change, a rate
far higher than is usually observed in the fossil record. 8ee the description of the e/periment on page 3'?.
8everal populations eventually stopped respondingE they reached a selectin plateau8 in which response
to selection stopped.
8election plateau is caused by natural selection and not by loss of genetic variation.
B/treme genotypes have low fitness.
+ow fitness is due to hitchhiking of deleterious genes and pleiotropy.
2fter the e/periment was stopped, the mean bristle declined showing that genetic variation was still
present and that genotypes with fewer bristles had higher fitness.
#he response to selection is based on both genetic variation in the original population and new mutations
that occur.
-atural selection often opposes artificial selection.
95 /IRECTIONAL SELECTION IN NATURAL POPULATIONS
2n attempt has been made to measure the strength of natural selection on quantitative traits.
&ne inde/ uses the mean and variance of a trait measured within a single generation before and after
selection has occurred.
B. g. the measurements are made on 9uveniles and then on those individuals that successfully
survive to adulthood and reproduce.
0 = mean phonotypeE V = variance
0b = mean phenotype before selection
0a = mean phenotype after selection
Vb = variance before selection
Va = variance after selection
V1 = phenotypic variance
0a60b
#he in&e3 f the intensit( f &irectinal selectin is i = 6666666666
>V1
BP2*1+B8 &: 8B+BC#!&- &- K2-#!#2#!VB CH2G2C#BG8, pages 3'( 63F).
8ee the e/amples of"
8tabili0ing selection" #he 5arwinQs finch 9eo)p%:a fort%) on 5aphne !sland. #he conflicting
selection pressures create stabili0ing selection that, on average, favors an intermediate bill si0e.
8tabili0ing selection" %irth weight in humans. 8ee page 3'(.
8tabili0ing selection" 2ntagonistic agents of selection. #he larvae of the goldenrod gall fly, its
parasitoids and predators. #aken together, these enemies imposed rather strong stabili0ing
selection. 8ee page 3'(.
Bvolution observed"
#he codling moth Cy%a pomone**a, diapause timing, page 3'(.
%lackcap songbird Sy*&%a atr%(ap%**a, new migratory behavior, page 3'A.
8oapberry bug ;aera haemato*oma, new food sources, page 3'A.
Heavy metal and plant adaptation, page 3F).
Cod and bighorn sheep, page 3F).
8#GB-,#H &: -2#KG2+ 8B+BC#!&-
#he strength of natural selection appears to be quite modest, although strong selection has often been
recorded.
8tabili0ing selection and diversifying selection appear to be about equally common.
#here is a tendency for the strength of selection due to variation in mating success and female fecundity
to be greater, on average, than that of selection due to differences in survival.
:HAT 'AINTAINS GENETIC VARIATION IN $UANTITATIVE CHARACTERS;
#here are high levels of variation in quantitative traits in both large and small populations.
2lleles that contribute to quantitative traits are probably seldom selectively neutral.
*any quantitative characters are sub9ect to fairly intense selection.
,enes contributing to quantitative traits have pleiotropic effects of survival and other fitness components.
,enes that affect quantitative characters may have other primary functions and only incidentally affect
quantitative trait.
Characters that in themselves may be neutral can be sub9ect to indirect selection because of the
pleiotropic effects of the underlying genes.
,ene flow between populations with optimal phenotypes can maintain variation in both populations, but
studies of captive populations in which gene flow does not take place show that these populations do not
differ substantially from wild populations.
*K#2#!&-68B+BC#!&- %2+2-CB HO1&#HB8!8
,ene flow between populations with optimal phenotypes can maintain variation in each population.
+evels of polygenic variation reflect a balance between the erosion of variation by stabili0ing selection
and the input of new variation by mutation.
#here is some doubt that variation by mutation can counter balance the strong stabili0ing selection that
acts on many traits.
*any data are consistent with a model of mutation$selection balance in which stabili0ing selection on a
character acts on mutations of large effect at a few loci, while much of the variation consists of small
pleiotropic effects of slightly deleterious mutations at many other loci that have a variety of other primary
functions.
CORRELATE/ EVOLUTION OF $UANTITATIVE TRAITS
Bvolutionary change in one character is often correlated with change in other features. !f changes by
selection occur in one trait, other traits will also change.
#he overall phenotype must be maintained in order to avoid deleterious changes.
C&GGB+2#B5 8B+BC#!&-
Characters that are functionally related are usually selected together.
,B-B#!C C&GGB+2#!&-
#he crrelatin cefficient 3r4 e/presses the magnitude or degree to which two traits are correlated.
#he correlation coefficient ranges from C1 for perfect correlation, when two features increase or decrease
together, to 61 when one feature increases e/actly in the same proportion as the other increasesE ) means
that the characters are not correlated at all.
#he evolution of a trait is governed both by selection on that trait and by selection on other traits with
which it is genetically correlated.
Phent(pic crrelatin 3r14 may have a genetic and an environmental component.
B. g difference in nutrition may affect two phenotypic traits of individuals with the same
genotype.
#his is referred as en"irn%ental crrelatin8 rE5
1henotypic correlation" r1 = r, C rB
Correlated variation may be caused by genetic differences that affect two characters, causing #enetic
crrelatin8 rG5
Causes of genetic correlation"
Lin1a#e &ise!uili)riu% among the genes that independently affect each character.
Pleitrp(, the influence of the same gene in two characters.
!n pleiotropy"
Perfect #enetic crrelatin occurs when one gene affects all characters in the same direction" all
increase or decrease" r,= 1.) or 61.)
I%perfect #enetic crrelatin occurs when one gene affects some characters in one direction 3C,
C or 6,64 while others have opposite effect 3C,6 or 6,C4, or affect only one of the characters.
,enetic correlation can change over time"
+inkage disequilibrium will decline with time due to recombination unless selection for the
adaptive gene combinations maintains it.
8ome alleles may become fi/ed so they contribute no variation and no correlation, while other
loci remain variable, and changes in frequency in these loci will affect r,.
-atural selection may affect %&ifier alleles that alter the pleiotropic effects of other loci.
8ee the e/ample of 2ustralian blowflies on page 3F3.
,enetic correlation among character and environmental correlation give rise to phenotypic correlation.
H&D ,B-B#!C C&GGB+2#!&- 2::BC#8 BV&+K#!&-
#he evolution of a trait is governed both by selection on that trait and by selection on other traits with
which it is genetically correlated.
,enetic correlation may be positive or negative.
,enetic correlations among characters can cause them to evolve in concert.
*ultiple characters may evolve as an integrated ensemble more rapidly if they are genetically correlated,
as when they are sub9ect to the same developmental controls.
B. g. characters that are functionally related like the organs of the body and the body si0e.
%ody si0e would evolve much more slowly in response to selection if every organ had to undergo
independent genetic change than if there were coordinated increases or decreases in the si0es of
the various organs.
5uring development the various organs grow in concert, and alleles that change body si0e have
correlated effects on most body parts.
,enetic correlation among characters may retard the rate of adaptive evolution, depending on
circumstances.
2 conflict may e/ist between the genetic correlation of characters and directional selection on those
characters.
Dhen such a conflict e/ists, the two characters may evolve to their optimal states only slowly, and may
evolve temporarily in a maladaptive direction.
B. g. selection for a deeper bill in the ,alapagos finch 9eo)p%:a fort%) caused average bill length
to increase, even though selection favored a shorter bill.
B.g. 2n e/ample from beef cattle breeding5 8election for fast growth is a common goal of
selection. %y doing so it is hard to avoid a change in the final adult weight. 2t the same time, the
selection will also result in larger calves being born, which can cause difficulties during calving if
the ratio between the weight of the calf and the cow is changed unfavorably.
http"$$www.kursus.kvl.dk$shares$vetgen$R1opgen$genetics$($'.htm
CAN GENETICS PRE/ICT LONG<TER' EVOLUTION;
8everal traits may be affected by the same set of genes, and selection on one of the traits would result in
changes in the others. #hese traits would be genetically correlated with one another. #he G<%atri3
describes these genetic correlations.
#he , matri/ is created by plotting the measurements of two traits, one in the /6a/is and the other on the
y6a/is.
!f large values for the trait in the y a/is was favored by natural selection, the values for the trait in
the / a/is will also increase as well over time, and individuals with large values for one trait will
also show large values for the other trait. #hese two traits are correlated and the dots will form an
oval in general outline. See F%'. 13.1<.
!f the plotted values are more or less evenly scattered and do not form an oval in general outline,
the traits are not correlated and evolved independently of one another.
http"$$www.bio.tamu.edu$users$a9ones$gmatri/online$whatisg.html
8ome characters may have very little variation and may constraint or determine the initial direction of
evolution.
8tudies have shown that genetic correlation between characters may remain consistent for a long time and
can influence the direction of selection. 8ome investigators have found that the genetic correlations
among certain characters are fairly similar among geographic populations of a species or among closely
related species.
&ther studies have found lower similarities showing that the strength of genetic correlations can evolve
rather rapidly.
NOR'S OF REACTION
#he nr% f reactin of a genotype is the set of phenotypes it e/presses in different environments.
L:or e&ery genotype, phenotypic trait, and environmental variable, a different norm of reaction can e/istE in other words, an
enormous comple/ity can e/ist in the interrelationships between genetic and environmental factors in determining traits.M
http"$$en.wikipedia.org$wiki$-ormsRofRreaction
PHENOTYPIC PLASTICITY
Phent(pic plasticit(" &rganisms with a given genotype may change its phenotype in order to adapt to
environmental changes.
2 single genotype may produce radically different phenotypes in response to environmental stimuli.
2daptive phenotypic plasticity has evolved by natural selection for those genotypes with norms of
reaction that most nearly yield the optimal phenotype for the various environments the organism
commonly encounters.
B.g. the leaf form of the aquatic plant ,anun(u*u) a-uat%*%) depends on whether it is submerged,
aerial or situated at the air6water interface during development.
#he following e/amples are from" http"$$en.wikipedia.org$wiki$1henotypicRplasticity
B. g. !nSsocial insects, colonies of which depend on the division of their members into distinct
castes, such as workers and guards. #hese two castes differ dramatically in appearance and
behavior. However, these differences are not geneticE they arise during development and depend
on the manner of treatment of the eggs by the queen and the workers, who manipulate such
factors as embryonic diet and incubation temperature. #he genome of each individual contains all
the instructions needed to develop into any one of several HmorphsH, but only the genes that form
part of one developmental program are activated.
B. g. !n epidemiology, a popular theory is that rising incidences of coronary heart disease and
#ype !! diabetes in human populations undergoing industriali0ation is due to a mismatch between
a metabolic phenotype determined in development and the nutritional environment an individual
is subsequently e/posed to. #his is known as the H#hrifty phenotypeH hypothesis.
CANALI=ATION
Canali7atin is the buffering of developmental pathways that tend to produce a standard phenotype
despite environmental fluctuations and underlying genetic variability" developmental homeostasis.
Kniform e/pression of traits or patterns of development despite slight changes in environment or
genotype.
*aintenance of a single adaptive phenotype.
2 trait is canali0ed when the same phenotype is produced in a wide range of genotypic and
environmental backgrounds.
Canali0ation occurs when selection allows a larger number of different genotypes to produce the same
phenotype.
8election for genotypes that can produce the same phenotype despite their variability.
Genetic assi%ilatin has taken place when a character state that initially developed in response to the
environment can become genetically determined and inherited.
Canali0ed characters include threshold characters, in which underlying polygenic variation is not
phenotypically e/pressed unless a drastic mutation or environmental perturbation breaks down
canali0ation.
#he following quotation refers to the e/periment performed by Daddington on the effect of heat shock
and the development of crossvainless trait in Dro)oph%*a on page 3F(,
L2s e/plained by D255!-,#&-, the development of crossveins and

other apparently very stable morphological traits can be
influenced

by environmental disturbances above a certain threshold of intensity,

but individuals from wild6type populations have
a threshold

so high that only an unusually strong stimulus, such as a heat

shock, can effectively induce a modified e/pression.
2ccording

to D255!-,#&-Hs e/planation, the phenotypic uniformity generally

observed in these traits can easily coe/ist with
the abundant

genetic variability demonstrated by artificial selection in

assimilation e/periments. 2lthough different genotypes
available

in a population are sensitive to different threshold values

of e/ternal stimuli, phenotypic variation does not arise if

all of
them have too high a threshold to be affected by the

disturbances prevailing in the usual environment. However, when

an
e/ceptionally severe disturbance occurs, the subpopulation

of individuals in which a phenotypic change is induced is necessarily

enriched for the most sensitive genotypes, which provide the

material for artificial selection.M
Canali7atin8 Genetic Assi%ilatin an& Prea&aptatin> A $uantitati"e Genetic '&el5
Ilan Eshel an& Carl 'atessi5 ,enetics, Vol. 1'A, ;11A6;133, 2ugust 1AA(
,enotypes of flies differ in their degree of susceptibility to the influence of the environment 3heat shock4
. they differ in their degree of canali0ation so that some are less easily deflected into an aberrant
developmental pattern.
8election for this pattern favors alleles that canali0e development into the newly favored pathway.
2s such alleles accumulate, less environmental stimulus is required to produce the new phenotype.
Daddington e/plained that genetic assimilation provided a potentially beneficial mechanism for
populations under stress
#he phenotype produced under stress becomes the phenotype for every condition.
LCanali0ation implies that a genotypeHs phenotype remains relatively invariant when individuals of a particular genotype are
e/posed to different environments 3environmental canali0ation4 or when individuals of the same single6 or multilocus genotype
differ in their genetic background 3genetic canali0ation4. Consequently, genetic canali0ation can be viewed as a particular kind of
epistasis, and environmental canali0ation and phenotypic plasticity are two aspects of the same phenomenon. Canali0ation results
in the accumulation of phenotypically cryptic genetic variation, which can be released after a Tdecanali0ingT event. #hus,
canali0ed genotypes maintain a cryptic potential for e/pressing particular phenotypes, which are only uncovered under particular
decanali0ing environmental or genetic conditions. 8election may then act on this newly released genetic variation. #he
accumulation of cryptic genetic variation by canali0ation may therefore increase evolvability at the population level by leading to
phenotypic diversification under decanali0ing conditions. &n the other hand, under canali0ing conditions, a ma9or part of the
segregating genetic variation may remain phenotypically crypticE canali0ation may therefore, at least temporarily, constrain
phenotypic evolution. *echanistically, canali0ation can be understood in terms of transmission patterns, such as epistasis,
pleiotropy, and genotype by environment interactions, and in terms of genetic redundancy, modularity, and emergent properties
of gene networks and biochemical pathways. Dhile different forms of selection can favor canali0ation, the requirements for its
evolution are typically rather restrictive. 2lthough there are several methods to detect canali0ation, there are still serious
problems with unambiguously demonstrating canali0ation, particularly its adaptive value.M The e"lutinar( #enetics f
canali7atin5 Gev %iol. ;))F 8epE ()334";(<631?.
PHENOTYPIC PLASTICITY
2daptive phenotypic plasticity has evolved when a genotype has the capacity to produce different
phenotypes suitable for different environmental conditions.
1henotypic plasticity includes rapidly reversible changes in morphology, physiology, and behavior, as
well as developmental switches that cannot be reversed during the organismQs lifetime.
#he genotypes produce the optimal phenotype for the kind of environment most commonly encountered
by the species.
Dest6Bberhard 3;))34 has proposed that phenotypic changes induced by a new environment may
sometimes prove adaptive, and may later be genetically assimilated.
:ew cases of adaptation initiated by phenotypic plasticity have been well documented so far.
EVOLUTION OF VARIA0ILITY
Variation refers to the differences actually present in a sample or species.
Variability refers to the potential to vary.
#he variability of individual characters is affected by the evolution of canali0ation.
2 character that is insensitive to alteration by environmental factors in environmentally canali0ed.
,enetic canali0ation describes insensitivity of a character to mutations.
!n genetic canali0ation, the phenotype remains unchanged when the genes underlying its development
vary.
LSgenetic canali0ation can be viewed as a particular kind of epistasis, and environmental canali0ation and phenotypic plasticity
are two aspects of the same phenomenon. Canali0ation results in the accumulation of phenotypically cryptic genetic variation,
which can be released after a Tdecanali0ingT event. #hus, canali0ed genotypes maintain a cryptic potential for e/pressing
particular phenotypes, which are only uncovered under particular decanali0ing environmental or genetic conditions. 8election
may then act on this newly released genetic variation.M The e"lutinar( #enetics f canali7atin5 Gev %iol. ()334";(<631?E
;))F 8ep.
Threshl& traits are controlled by polygenic variation rather than by single loci.
#he polygenic variation is not e/pressed phenotypically unless development is disturbed by substantially
beyond the threshold by a large enough genetic or environmental change.
#he evolution of canali0ation may e/plain the constancy of some characters over vast periods of
evolutionary time.
B. g. Barly 5evonian amphibians had a variable number of eight or nine toesE then five6toed
limbs evolved and became canali0ed, so vertebrate tetrapods since then have had more than five
digits.
#he h(pthesis f %rphl#ical r phent(pic inte#ratin holds that functionally related
characteristics should be genetically correlated with one another.
Dagner and 2ltenberg 31AA?4 have shown theoretically that prolonged directional selection favors
modifier alleles that enhance a pleiotropic correlation between functionally related traits.
B.g. if it were functionally important for the upper and lower mandibles of a birdQs bill to be the
same length, then selection for a longer bill would include selection for alleles that coordinate the
development of the two mandibles, creating a pleiotropic correlation between them.
:or additional information see" http"$$eebweb.ari0ona.edu$faculty$badyaev$papers$<3.pdf
GENETIC CONSTRAINTS ON EVOLUTION
*any species have become e/tinct because of their failure to adapt. Dhy didnQt they adaptN
Bvolution may be constrained by the amount of genetic variation 3V24 in individual characters and genetic
correlation 3r,4 among characters
8ome adaptations are the result of a single mutation rather than great genetic variation, e.g. resistance of
Cu*e6 p%p%en) to organophosphate insecticides is due to a single mutation that spread around the world by
gene flow.
Gecent adaptive evolution has been based on genetic variation or new mutations.