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UNIT
- THE PROCESS OF
EVOLUTIONARY CHANGE
Structure
11.1 Introduction
Objectives
11.2 The Basis for Natural Selection
Prodigality of Nature
Factors that Limit Reproductive Potential
Variability in Populations
Natural Selection
11.3 The Sources and Expression of Variability
Sources of Vatiability
Expression of Variability
11.4 Concept of Fitness
11.5 Natural Selection Under Different Environmental Conditions
Action of Natural Selection U i ~ d e rUniform Environments
Action of Natural Selection in Changing Environments
Action of Natural Selection if?Heterogenous Environments
11.6 Summary
11.7 Terminal Questions
11.8 Answers

1 11.1 INTRODUCTION
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Units 9 and 10 of this Block introduced you to the concept of Evolutionary Biology.
The Unit 9 dealt with the origin of the concept of evolution and the major
contributors t o the dhrelopment of evolutionary thought. In that Unit you also learnt
about Darwin, who based on his extensive observations and interpretations showed
that evolution is a simple fact of nature. It was Theodosius Dobzhansky, a population
geneticist and one of the trinity to structure the modern synthetic theory of Evolution,
who emphatically stated, "Nothing in Biology makes sense, except in the light of
evolution". Today the Darwinian concept of evolution through natural selection has
stood the test of time. As Julian Huxley rightly pointed out "Darwin rendered
evolution inescapable a,s a fact, comprehensible as a process and all embracing as a
concept". In Unit 10 of this Block you were cited several examples from studies on
palaeontology , biogeography, anatomy, embryology, physiology and biochemistry as
evidence in support of the evolutionary process. In this Unit we will more specifically
discuss the mechanism of evolutionary change. We will elaborate on the Darwinian
concept and look into its basic tenets. There will be a detailed discussion on the
sources of variability, since variability is the raw material on which natural selection
acts to produce adaptations. We will also explain the concept of Darwinian fitness
and relate it to the reproductive success of organisms. Finally you will also become
familiar with the different ways in which selection can act. In other words, this Unit
will highlight that natural selection promotes structural, functional and ecological
adaptations of individuals, populations and species to their existing environments.

Objectives
After studying this unit you should be able to:
justify the concept that natural selection is fundamentally a process of differential
reproduction,
identify the sources of variability which furnish material on which selection acts,
define the term fitness or adaptive value and explain how it is a measure of
selective process, and
examine the working of natural selection in homogenous, heterogenous and
changing environments.
 11.2 THE BASISTOR
NATURAL SELECTION
In the first Unit of this Block, you have learnt the basis on which Darwin propounded his
thesis of evolution as descent with modification through variation and natural
selection.
Darwin put fort11the notion of the survival of the fittest. How do we define 'fittest'?
The answer usually given is those who survive. So you can readily see that this is a
circular agrument and doesn't really help us understand what really happens
following natural selection quite clearly. The idea of fitness as a matter of survival
will not do. Therefore in the course of the development of what we have earlier
referred to aS the modern synthesis the notion of fitness was related to the ability of
the individual to maximise the number of progeny left in the next generation. In other
words the notion of fitness was translated into reproductive success. Given the idea
of fitness as being measurable in terms of reproductive success we can now go back
to basic tenets of Darwinism detailed in Unit 9. The first of the five observations
mentioned therein was the tendency on the part of living organisms to reproduce: In
an exponential manner. We noted that the consequence of this is competition for
available resources which results in "the survival of the fittest" Clearly, therefore.
in3terms of reproductive ability nature is truly prodigal.

11.2.1 The Prodigality of Nature

By prodigality of nature it is meant that organisms have an enormous potential to
reproduce. A carp or salmon is known to lay over a million eggs. A frog may lay Bs
many as 12,000 eggs. If all the eggs survive and reproduce, then within a short time
the entire earth would be strewn with frogs. Darwin himself worked out the
reprodilcti-Werates of elephants which are known as slowest breeders of all known
animals. Assuming an elephant during its 100 years of existence gives bjrth to six
calves in sixty years of its active reproductive life, in around 750 years there should
be at least 19 million elephants living in the world. Several such examples could be
quoted to highlight the fact that organisms have an immense potential for
reproduction.
How is it that in spite of an enormous potential to reproduce, the numbers of most
of the species are always maintained at optimal levels? How is it that lakes are not
choked with fishes, or fields not strewn with frogs or the earth not crowded with
elephants? This is because that various limiting factors in the environment, both
biotic and abiotic, keep a check on the increase in numbers.

11.2.2 Factors that Limit the Reproductive Potential

9 Organisms are made to compete for their needs from the environment. The
competition as we pointed earlier could be for the food and territory, to overcome
caving CaPacIb' of environment iKI the adverse climatic conditions, to escape from predators or tp combat an infectious
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10
diskase. This is indeed the "struggle" It is this "struggb" that keeps the population
u numbers under manageable levels and from increasing on a logrithmic scale. We
5
.- earlier referred to the reproductive potential of frogs. If the potential is fully realised,
o
c
"- then in a short time, they would be placing a severe strain on the resources on which
0 ,
they depend and soon all of them would starve. However, this is not the case. Most
8 of the population sizes do not increase enormously but they only fluctuate to asmall
5
z
extent from time to time. Some other populations'however, do increase aramatically
in certain seasons and decrease even more dramatically in certain other seasons. In
laboratory one can grow cell cultures in which any one of the resources such as food,
0 Time or generations space etc. is limited. Under these circumstan~esthe population growth rate of the
cells which is exponential to begin with decreases and levels off: The resulting sigmoid
,Fig. u.1:me w p h #howlng the curve is characteristic of biological growth in general. (Fig. 11.1) The factors that we
slgmoid growth curve of popuhtfon. mentioned earlier are the ones which delimit the numbers or size to the carrying
capacity of the environment in natural populations. For instance, the 'carrying
capacity of the environment for plants is controlled by the ambunt of spact! required
'by the individuals of their own and other species. No new seeds or shoots can develop
in that particular area until the older plants fall. Population sizes in animals, as
mdntioned earlier can be limited by food supply. Here the number of individuals ,
increases until the food shortage occurs. Such a shortage of resources would decrease,
.46 the reproduction. Now a situation would arise where the population size is decreased '
 and the carrying capacity of the environment becomes high enough to support more
number of individ,uals. The cycle thus repeats itself.
he- ol ~vdutlollnrychnnge

By word of caution we may mention here that the term "struggle" should not be If a small number of individuals is
interpreted too literally. In fact after Darwin published his, 'Origin of Species', there introduced into a suitable
environment with abundant
was criticism that Darwin's theory suggested war, aggression, fighting between races resources, birth rates are high and
and classes, and ethnic competitions. But as rightly pointed out by G.G. Simpson death rates are low. As more
such a "mature red in tooth and claw" approach to natural selection is unfortunate individuals are added to
The struggle as mentioned by Darwin is a verysubtle one. For instance, plants which population, resources become
'struggle' in dry environments develop devices which would prevent ioss of water scarce and death rate increases.
Also the birth rate decreases. The
from them and not by sucking water from each other. growth of the population stops as
the density reaches the maximum
So from among a population of individuals, if only a certain proportion is able to number of individuals that
overcome the 'struggle' and survive to produce the next generation's offspring, the environment can support. This
density is known as canying
questions that come to one's mind are: Is it that chance aldne determines which of
capacity.
the individuals should survive and which ones should die? If not, what is it .that
determines the survival of individuals? These questions lead us to the next element
in the process of natural selection, namely, the element of variability in populations.

11.2.3 Variability in Populations

Variability refers to the differences in heritable traits exhibited by the individuals of
a species. One of the major contributions of Darwin to the study of evolution was his
book of 1868, 'The Variation of Animals and Plants under Doxnestication'. Danyin
pointed out that populations of living organisms are not composed of identical
individuals.

It is indeed true that even monozygotic twins may express differences between them.
Onecan cite a number of examples to indicate the differences in different traits in a
population. Certain traits are continuously varying such as the height or colour, and
certain other traits exhibit two or a few distinct, different categories of individuals
1 with no intermediates. For example in human population, there are P'C tasters and
non-tasters; individuals with blood groups A or B or AB or 0 ,individuals with black
or brown or blue eyes, individuals suffering from sickle cell anaemia and individuals
who are normal, so on and so forth. Sweet peas may have either red flowers or white
ones; Drosophila may have normal or vestigial wings, red or white eyes etc.
Essentially the major source of variability in organisms is the genetic variability
caused by mutations of all kinds (Refer to Unit 16 of LSE-04 genetics) and genetic
recombination. Also the subtle effects of environment add another dimension to the
I1 variability of traits in natural populations. An oft cited example is the differences
exhibited in a number of traits of monozygotic quadruplets of a nine-banded
armadillo. The monozygotic offspring could exhibit such differences, because of the
differences in the uterine environment in which they undergo development. It should
be obvious to you that four genetically different zygotes developing in the same uterus
should be even more different, We will have a more detailed discussion on the role
of natural selection in a changing environment later in this unit.

11.2.4 Natural Selection

Assuming that variability of populations were non-genetic, that is, not controlled by
genetic material, once again chance events alone would determine which of the
organisms would survive. But phenotypic traits of a 1 organisms'have a genetic
component and heritable to some degree. If an individual has some form of a trait
that gives him a better chance of survival in a given environment, evidently his
offspring inherit the trait. This is known as differential survival. Differential survival
*auld refer to the survival of individuals in an environment to which they are well
adapted. Individuals who do not possess such adaptations for the given environment
may not survive to reproduce. The adaptations are genetically controlled and
heritable. Differential survival leads to differential reproduction of alleles responsible
for the su~vivalof the next generation. In fact it is customary to say that natural
selection is synonymous with differential reproduction. The consequence of
differential reproduction is not so much that individuals survive, but that such
individuals are able to perpetuate their genes in subsequent generations. Thus what
really survives is the set of genes.
Natural selection can therefore mean differential reproduction among members of
the sarne gene pool. We earlier said that selection would promote those alleles that
give rise to phenotypes that are better adapted to a given environment. In other
words, the frequencies of 'such alleles would tend to increase and those of ill-adapted
ones would tend to decrease. What we are sayingis that natural selection is a matter
of changes in gene frequencies (see box for a brief discussion on changes in gene
frequencies). Differential reproduction has a dual role to perform. One it leads to or
maintains adaptations and therefore determines whether individuals of a population
or species will survive; and two, it is a process which determines what changes will
occur in the frequencies of alleles in a gene pool of sexually reproducing individuals.

GENE FREQUENCIES AND THE CHANGES THEY 'UNDERGO IN

POPULATIONS

G.H. Hardy, an English mathematician and W. Weinberg, a German physician

independently formulated the central theorem of population genetics.
According to them, the frequencies of a pair o i alleles in a population would
remain constant generation after generation, provided that the population is
infinitely large and randomly mating one, and that there is no mutation, no
selection, no recombination and no gene migration in the population. But in
natural populations, these conditions can not be met and therefore, the
equilibrium conditions can not be obtained. In other words, in natural
populations gene frequencies cannot be expected to remain constant in every
generation, since phenomena such as mutation, recombination, migration etc.;
occur randomly. Let us try to explain how gene frequencies change in
population with a simple example.

Assurne that two alleles A and a, have frequency p and q respectively in a large
population. Let us assume that frequency of A and that of a put together is
equal to 1. Then, (p + q) = 1 or (1 - p) = q. Let us say that the two alleles
A and a form three genotypes, AA, Aa and aa. The frequency of the genotypes
+
would be the binomial expansion of (p q); that is (p + q)2
(p + q)2 = P2 + 2pq q2 +
Therefore, the frequency of AA
frequency of aa = q2.
- p2, the frequency of Aa = 2pq and the

+
Since we said (p + q) = 1, (p q)2 is also equal to 1. The law states that at.
equilibrium conditions, the frequencies will not change. Therefore, in next
generation, frequency of A would be determined by the number of gametes
carrying A, and this would depend on the number of carriers of A in the
population. Then frequency of A = p2 + pq (one half of 2pq) = p' + p (1 - p) =
P2 + p - p2 = p and frequency of a = (1 - p) = q. Then in the absence of any
force tending to change gene frequency - mutation, selection, gene flow etc., the
frequency remains constant from generation to generation.

But evolution manifests as change in gene trequencies. Let us briefly look into
the effect of mutation on gene frequencies. Let us say that allele A mutates to
a at a rate of u. Assuming the frequency of A is p and that of a is q initially,
then in the next generation frequency of A would be:

PI = Po - Po"
po is the initial frequency, pl is the frequency in the next generation and pou is
the frequency of mutated alleles.

In other words, the frequency of A has changed from p to po (1 - u) and that

of a will be [I - (po (1 - u ) ] or (1 - p,). A more detailed discussion on the
subject can be obtained from the unit on 'Behaviour of Genes in Population' in
LSE-03 Genetics course.
 On this note we end this discussion on the basic Darwinian principles. You may
attempt the fol!owing SAQs before we look into the possible sources of chromosomal
and gnetic variability and cite certain examples to illustrate the expression of
variability.

SAQ 1
Say whethen the foilswing statements are true or false:

i) Darwin and Wallacc were the coauthors of the book 'Ollgin of

of Species' ( )

ii) It was Dobzhansky who said that 'nothing in biology makes sense
except in the light of evolution'. ( )

iii) Variations are rare in nature. ( )

I
iv) By 'struggle for existence', Darwin meant fighting among the
individuals of a species with weapons. ( )

V) One way to describe natural selection is that it refers to changes

in gene frequencies in a population. ( 1
v) Adaptations evolve because of the interaction between the
genotype and environment. ( )

11.3 THE SOURCES AND EXPRESSION OF

VARIABILITY
In the preceding section we emphasised the fact that cvolution or the continuous
, adaptations to the environment can take place only when pheiiotypes exhibit
variability. In the absense of variability, the chances arc that existing living organisms
would become extinct since with the change in the environment these organisms
would die without leaving progeny. Fortunatcly in living systems mutations and
recombinations generate the required variability. However, organisms often strive to
arrive at populations which are more or less uniform. This is possible because
selection process acts in such a way that a certain degree of uniformity of phenotype
is imposed among individuals of n population. If an individual were to be vcry
different in some way from thc rest of the population. he would be less well adapted
to the environment as compared to other members of the population and therefore
I unable to compete successfully with them. However, in a continuously changing
environment, a slight phenotypic variability should be maintained in a population. In
a changing environment no phenotype remains optimally adapted for long. This is
the precise reason why variability needs to be generated continuously in a population
and used to develop adaptations in the way Darwin suggested. We shall look into the
mechanisms that generate variability.

: 11.3.1 Sources of Variability

I
Mutations and genetic recombination (in sexually reproducing organisms) are major
! sources of variations in natural populations, In this section we will briefly mention
I the various types of mutations, their rates and how they are caused. For a more
detailed account of mutations and mutagenesis you may refer to Unit 16 of Block 3
! of the LSE-03 Genetics course.

Mutations
1 The term mutation is used to designate the process by which changes arise in the
iI genetic material and, the end products of such processes. Mritations form a separate
1 category of sources of variability, distinct' from variability arising from genetic
recombination o r independent assortment of chromosomes, characteristic of sexually
reproducing individuals. There are two categories of mutations. '(1) Chromosomal

i mutations which affect the'number of chromosomes and the number or the

arrangement of gencs in a chromosome; (2) gene mutations which affect only one or
a few nucleotides in a gcne. We shall now tiiscuss these two aspects of n~uialion~.

1. Chromosomal mutations:
Following are the ways in which chromosomal mutations occur: For a detailed
discussion on chron~osomalmutations refer to Units 9 and 1 0 of LSE-03 t:ienetics
Course.
a) Changes in number of genes in a ehronaosorne.
i) Deficiency or rleletion : Refers to the loss of a segment of a chromosome
containing one or several genes.
ii) Duplication : Refers to the occurrence of rriore than once of one or more genes
in a chromosome. Duplications often occur in random, that is, two or more
duplicated segments may lie adjacent to each other on the same chromosome.
b) Changes in the arrangement of genes in n chrornosome
i) bzversion : The givela sequence within a segment of a chroinosome is reverted
ii) Trdnslocation : The location of a block of genes is changed in the chromosome.
Usually translocation is reciprocal which mearns that it is an exchange of a block
of genes between two non-homologous chromosomes.
C) Changes in chromosome number
i) Aneuploidy : One or inore chron~osomesof a normal set nlay bc lacking or
present in excess. The term riullison~yrefers to absence of both the chromosomes
of a pair. Monosomy, trisomy, tetrasomy etc., refer to the occurrence of a given
chromosome once, thrice, four times ctc., resycctively in a diploid organism.
ii) Polyploidy : Refers to the occurrence of more than two sets of chromosomes in
an individual. Most organisms are diploid, that is they have two sets of
chromosomes in their somatic cells, but only one in their gametic cells. Polyploid
organisms would be triploid when they have three sets of chromosomes,
tetraploid if they have four sets and so on. Polyploidy occurs frequently in plants
but is a rare phenomenon in animals. Polyploidy has been a common means of
speciation in plants. Many of thc cultivated plants, such as wheat, oats, tobacco,
potato, banana, coffee, sugarcane and many flowering plants are polyploids.
Plant breeders always attempt to induce polyploidy by artificial means to create
new varieties. Polyploidy can be regarded as thc only means in bisexual
organisms by which a new species could arise in one step.
2. Gene mutatiorns
Gene or point mutations occur when the DNA sequence of a gene is altered and the
new nucleotide sequence is passed on to the offspring. This occurs either due to
addition or deletion or substitution of one or a few nucleoticles. Nucleotide
substitutions can be either transitions or transversions. Transitions are replacements
of a purine by another purine (A by G or vice versa) or a pyrimidiile by another
pyrimidine (C by T or vice versa). Transversions are replacements of a purine by the
pyrimidine or vice versa (G or A by C or T or vice versa). We will briefly explain
the different types of gene mutations.
a) Substitutions : substitution of one base by another would result in an altered
amino acid in a polypeptide chain. For instance triplet AAT in DNA (UUA in
mRNA) would specify leucine. But if the first A is ieplaced by C, it will code for
valine. Some of the codons are degenerate and substitutions may not alter the amino
acid specified. For instance, in the triplet AAT if the first A is replaced by G the
amino acid coded would still be leucine. Gene mutations which do not normally affect
the active site of a protein will not alter its biological functions. But nucleotide
substitutions that change a triplet coding lor an amino acid into a termination codon
would produce adverse effects. Once again in AAT, if the second A is replaced hy
T ( A n ) , the resulting rnRNA codon IJAA is a termination codon. When a
termination codon is present in the middle of an mRNA molecule the subsequent
codons are not translated and an incomplete polypeptide will be released from
ribosomes.
b) Additions and deletions : Addition or deletion of a nucleotide pair in the DNA
sequence of a structural gene often results in an altered sequence of amino acids in
the coded polypeptide. Essentially such additions or deletions shift the reading frame
 of nucleotide seqsencg from the point of addition or deletion to the end of the
molecule. Look ai'rhe following DNA sequence.
CAT - CAT - C A T - C A T - C A T -
t
If a nucleotide T were to be inserted immediately after the first C; the sequence would
read as
- = T A- TCA - T C A - T C A - TCA - T'
The original DNA sequence when ,transcribed and translated would yield five
successive valine residues. But the altered sequence would correspondingly read one
aspartate and four serine residues. Mutations resulting from the addition or deletion
of base pairs are called frameshift mutations.
So far we have discussed the several ways in which the genetic material could be
altered. Mutations could be either spontaneous or induced. In recent years there has
been a more clear understanding of the causes of mutational process. Tautomeric
shifts, ultraviolet radiations, chemicals such as hydroxylarnine, nitrous acid and a
variety of alkylating agents can cause permanent alterations of nitrogenous bases.
Aromatic compounds such as acridines can cause addition o r deletion of one to
more than twenty bases in DNA.
We have briefly discussed mutations, a way or source of variability in natural
populations. Together with recombinations occurring'at the tjnie of meiosis they form
the raw materials for selection to act on. Before we discuss briefly about
recombination, let us have a quick look at mutation rates in organisms. For a detailed
discussion on mutation rates you may refer to the Unit 16 of Block 3 of LSE-03
Genetics course.

3. Mutation rates
Each gene has a characteristic mutation rate. Mutation rates are often described as
an average per gene basis. For instance, Drosophila has one detectable mutation per
10,000 loci, that is, 0.01% per locus. In humans spontaneous mutation rates have
been calculated for some of the dominant lethi1 genes such as retinablastoma,
chondrodystrophy and Huntington's chorea. These genes have rates varying from 0.01%
to 0.001%per locus which means that one out of 10,000 or 100,000 loci mutates. This
, is true of most of eukaryotic mutation rates which range from to per locus.
-'
Jn prokaryotes the rates may be much lower, ranging from 10 to 10-"'per locus.
Mutations can also occur in reverse direction by which it is ineant that a mutant gene
I
can mutate back to its wild type form. However, reverse or back mutation rates are
much lower than the forward mutation rates.
I 4. Genetic recombination
Mutations are the only kind of variability generating mechanism in prokaryotes and
asexually reproducing organisms. But in eukaryotes where sexual recombination
occurs, even in one generation there is a marked reshuffling of genes in the
chromosomes which amplifies the genetic variability in a population. The kind of
variability generated by recombination is tested by natural selection in as many
genetic backgrounds as possible and in as short time as possible (say in one
generation). Thus one of the greatest and fundamental advantages of sexual
reproduction is the generation of variability. Provided the populations are large, even
if mutations do not exist, the mechanism of sexual recombination alone would be
, generating new genotypes for very long time.

, I
11.3.2 Expression of Variability
In the previous sub-section we discussed the possible ways by which variabiliti can
, be generated. We shall now examine one instance that illustrates the consequence of
variability on phenotype.
1

i Any genotype is essentially an encoded hereditary information for a specific

/ phenotype over a small range of environmental conditions. In other words a
: phenotype which arises as an interaction between genotype and the environment is
i represented by a structure o r function of an organism that would possibIy come under
1 the influence of natural selection. Let us now look into an instructive example of

; sickle cell anaemia - a genetic disease, the allele for which manifests differently in
different environments under the influence of natural selection.
I
 Sickle cell anaemia in man is caused by a defective chain of haemoglobin. The
abnormal haemoglobin known as MbS differs from natural haemoglobin (HhA) in
one amino acid. A substitutional mutation causes the replacement of glutamic acid
by valine in the sixth position from the amino terminal end of the Q chain polypeptide.
This is because instead of the triplet GAA or GAG which codes for glutarnic acid in
normal chains, the abnormal mRNA has GUA or GUG which codes for valine. IdbS
in deoxygenated state precipitates out of solution forming elongate crystals. 'The
formation of crystals in turn produces a characteristic deformation of RBCs and the
cells acquire a crescent or sickle shape, hence the name sickle cell anaemia. The
deformation of red cell weakens the red cell membrane and lyses the cells. The loss
of red cells and haemoglobin leads to anaemia and this causes oxygen deficiency to
the tissues per unit time. Pt, single substitution, more specifically, replacing the
thymine by adenine in the second position of the sixth triplet of the P chain gene
triggers a variety of responses that affects several aspects of the pfnysiology of the
organism as shown in Fig. 11.2. Individuals who are homozygous (HbSiHbS) for the
gene usually die early in life. In heterozygotes ( W b ~ M b ssome
) proporrion of red
cells lyse periodically but this does not cause any serious harm to the individual and
in any case the survival is not affected.
-. ,

Tl~yi~iinc.
replaced by adenine )
, ,in beta chain
.. . . -. .

i
I . .. . 9

- -----..----c~..__.__
Abnormal globin .!

Abnormal hemoglobin j
.- ~. ,I

I
,

. - . . . . .
; Sickling of
j r;d blood ct.113 1 '
.
.: L-
I
Clumj7ing of cells itlrd
cloggjng of circulalioa~

fhpid destruction
1
Local failures i c ~
of sickle cells blood supply

Ovcritctivi~y'11
b n e marrow

Wcaklress and Urain

lassitude damage hluscle
join1 and
damage damage
I-ung Kidney
dani4rge

Skrle~al Jmpaircd nirntal Poor physical

changes f unctiunin~ development Paralysis Rheumdtiam Pnrunlvnia
failure

Fig. 11.2 : Effects of sicye ceU anaemie. Thc figure shows that a substitution of a dtrogenous b a ~ eby
another, triggers a variety of responses that nearly afTectseveral aspectsol organism's physlalogy.

In other mammals sickle cells do appear in population, but do not lead to the lysis
of erythrocytes. For instance, deer populations have been shown to possess sickle
cells extensively in their blood. It is suggested that the thickness of the red cell
membranes of deer populations would prevent the lysis of the cells. What is more
interesting is that the abnormal allele of the p chain gene is present in high frequency
in some human populations. How can a gene that is imposing deleterious andlethal
effects on its homozygotes be present in such large numbers in certain human
populations? This essentially means that natural selection in some places favours or
at least is not eliminating the allele. We earlier said that in heterozygotes there is 1

only a small loss of haemoglobin. But all other factors being equal, even such a small
 .ass cannot be considered as a favourable condition and natural selection would The P m m s of Evdutiowy Change
eliminate the allele from the population. Under these circumstances there should be
some compelling reasons for the maintenarm of the _allelein large numbers in A e c a n
and cenain Asian populations. The allele for sickle cell haemoglobin (HbS) almost
coexists with normal allele (HbA).
Evidence is available that hetcrozygotes canying HbS are resistant to falciparum
malaria. In the malarial belt of Africa and Asia, the frequency of HbS allele is quite
high. Medical examination of children in hospitals has indicated that those with sickle
cell trait (not the disease) d o not exhibit falciparurn malaria symptoms. Studies have
shown that even when bitten by carrier mosquitoes, individuals do not contract the
malaria provided they are heterozygous to the haemoglobin p chain gene, whereas
normal homozygotes ( ~ ~ did contract
A ) the disease. The HbA/HbS
heterozygote has a greater advantage than the normal haemoglobin homozygote
(HbAIHbA) in that the former gets a complete protection against the malarial attack.
Sickle cell anaemia is an example, where having a defective allele or part of the
genotype proves to be an adaptation.
When genes are represented by only a single wild type allele obviously their
variability will be low in the population. Certain genes may exhibit greater variability.
Physiological traits appear to have a wide range of variability as against anatomical
traits. The coefficient of variation (a measure of variability- see marginal remarks)
for a randomly chosen set of physiological traits in animals ranges from 7.1 to 304.
Anatomical traits in mammals have a range from 4 to 10. It is also true that variability
of certain of the traits is indirectly related to some final important physiological
function. For instance blood pressure is a physiological trait closely monitored by
natural selection showing a low variability among the individuals of a population
under identical conditions. Contrarily the heart rate, capillary diameter, or the force
of ventricular contraction could be highly variable in a population. Each of these
contributes to the final important product namely the heart beat. The blood pressure
may be arrived almost to a fixed value by different contributory values of various .
components. For example, it could be said that the value 6 can be atrived at by 12/2,
18/3, 24/4 etc.
Apparently there is not much of a variation in anatomical traits such as the length of
head or relative length of forelimbs or in certain physiological traits such as stamina,
swimming speed or blood pressure etc. Once developed these traits appear more o r
less fixed. Such constancy is seen in the structure of protein molecules as well. For
instance, analysis of amino acid composition of the cytochrome C of 33 species of
diversely related organisms ranging from humans to sesame seeds has shown several
similarities. For instance amino acid positions 14, 17, 18 and 80 are invariant since
they all interact with heme molecule. Any change in these amino acids would
adversely affect the functioning of vital heme group which undergoes oxidation-
reduction reaction. Amino acids 80 to 85 are always hydrophobic, 99 to 104.i~ never
hydrophobic, 86 to 93 is charged and 94 to 98 is never charged. These charge
consistencies suggest that all these cyt&brome C molecules are folded in the same
way. In order to preserve the three dimensional structure of the molecllle,
selection has preserved the charge patteni of this vital protein.
.
. natural
W e end our discussion on sources and expression of variability here and b e h r e we,
move on to the next section, attend to the following SAQs.

SAQ 2
FilI in the blanks:
i) Continuous adaptations to the environment can take place only when
phenotypes exhibit .......................
ii) ~'h'emajorsources of variability inbnatural populations are ..................... and
.............................................
iii) Changes whichresult in the arrangement of genes in a chromosome are
......................and .......................
..
iv) ................... has been responsible in a major way for the evolution of
cultivated plants.
v) Mutations resulting fforn the addition or deletion of base pairs are called
......................
 vi) In sickle cell anaemia in P chain of haemoglobin ......................is substituted
for ....................... -
, vii) In African populations resistance to nlalaria is provided by
......................

%-

11.4 CONCEPT OF FITNESS --

While discussing the Darwinian premise of natural selection we observed that the
term selection is synonymous with non-random reproduction, and that the success of
the survivors is related to the number of offsprings they leave behind. Is it possible
to quantify such success? In other words can natural selection be quantified? The
answer to these questions is yes and is related to the notion of 'fitness'. The term
fitness variously known as Darwidan fitness or selective value or adaptive value is a
measure of the reproductive efficiencyof one genotype relative to another genotype
within a given population of species Let us now look into the details of the concept
of fitness.
Lei us .take an example to explain this concept.
Let alleles A and a form three genotype AA, Aa and aa. Let genotypes AA and Aa
produce four progeny each and genotype aa produce only two progeny. The fitness
or the adaptive value of genotypes AA and Aa would then be 414 = 1 and that of aa
would be 214 = 0.5. The two genotypes AA and Aa have similar fitness values and
aa has 50% uf the fitness value of the other two. The genotype with highest
reproductive efficiency is given the fitness value of 1 and the fitness of the other
genotypes are calculated relative to the one with maxinlum fitness value. Fitness is
denoted by the letter w. 'Table 11.1 explains to you the method of calculating the
fitness value for the various genotypes.
Table 11.1 : TabCe llPustretlag WARM a$ coleukutlon of adaptive value.ad selection
mfilcieni of genotypes
.. ---
Genotypes
81) Bb bb TOM

1. No, of genotypes in one 30 60 IS 105

generation
2. No. of zygotes produced in '90 NO 10 180
2nd generation
3. Average number of progeny 9W30 = 3 80160 6 1.33 10115 = O.ti6
4. Darwinian fitness (w) 313 = 1 1.3313= 0.44 0,6613 = 0.22
(relative reproductive
efficiency)
5. Selection coefficients = (1-w) 0 0.56 0.78

The measure of selection pressure on any genotype is referred to as selection

coeficient. Darwinian fitness and selection coefficient are related to each other by
the equation w = (1 - s) or s = (1 - w) where w is the Darwinian fitness and s is
the selection coefficient. s actually measures the amount of reduction in the fitness
of any genotype. Table 11.1 shows that the fitness of genotype BB to be one. This
means that the selectiorl coefficient for this genotype is zero which means that
no individual with this genotype is eliminated by the action of natural
se'lection. For genotype Bb and bb the selection coefficient would be 1-0.44 = 0.56
. and 1-0.22 = 0.78 respectively. In these cases 56% and 78% of the progeny are
elirnillated by natural selection. Fitness values arc useful in predicting the changes in
the frequency of genotypes from generation to generation. (]For a detailed discussion
on the allelic and genotypic frequencies and their changes in population refer to the
-
unit 20 Block 4 of LSE-03 Genetics course.)
From the above discussion it should be clear to you that the fitness of genotypes is
measured relative to each other and as such do not give us absolute values. It is the
relative reproductive efficiency of the different genotypes withiri a population that is
related to changes in gene frequencies and .this is essentially natural selection in
quantitative terms. Fitness values are affected by a number of factors all of which are
collectively referred t o as fitness components. These could be viability, role of The ~mcessd Evdutlmnry Chnnge
development, successful mating, fertility etc. The difference in fitness values which
different genotypes exhibit are due to the differences in one or several components.
For example thalassemia major, an inherited disease is caused by a defective cx chain I

locus. The affected individuals would die young. Therefore fitness values of such
individuals is zero. O n the other hand acbondroglastic dwarfs reproduce only 20%
as efficiently as rlormal individuals. In thalassernia individuals zero adaptive value is
due to the rlon-viability of individuals. In achondroplastic dwarfs it is due to reduced
fertility.

11.5 NATURAL SELECTION UNDER DIFFERENT

ENVIRBNbfENTAk CONDHTIONS
In this section we will be learning the action of natural selection on the populations
under different erivironmental conditions namely (1) environments which are more
or less uniform (2) environmznts which are constantly changing (3) environnlents
3
which are heterogeneous. $m
11.5.1 Actiorr sf Natural Selection in Uniform Environments Ir"
In the absence of any large scale envirorrmental char~gu,that is, in more or less Phenotype
uniform environments, populations often maintain a stable genetic constitution with
respect to many traits. Natural selection in the absence of environmental change Stabilizing
maintains a genetic homeostasis. This phenome~~on is known as normidisingselection. Selection

Several of the phenotypic, traits of individuals can be arranged on a linear scale. The
distribution curve of the traits usually takes a bell shape, so that the number of Before
individuals is greater at intermediate values and gradually decrease towards the Selection
extremes as shown in Fig. 11. Ja. As stated earlier, normalising selection occurs when
individuals with intermediate phenotypes are favoured and those wit11 extremes are
under selection pressure (11. Jb) . This tendency continues generation after
generation. If there is a strong selection pressure against the phenotypes occupying
the extremes of normal curve, then the population may show less variability although
the mean remains the sarne (Fig. 11.3~).Natural selection has more often a
normalising or stabilising effect on populations with mid-values for the traits and
individuals with intermediate values for their traits have better chances of survival.
For instance, new born infarits which weigh very less or very more than the average
weight have high rate of mortality. Contrarily infants of intermediate weight have less After
problems of survival. We discuss two examples below, one from nature and the other
from the experiments of Dobzhansky and Spassky to explain the concept of
normalising selection.

An interesting observation made by an American biologist H.C. Bumpus (1899)

/
Fig. 11.3 : Normallsing selection
provides a good explanation for normalising selection. Bumpus collected some 136 a) Before selection the popvlation
has a normal distribution or
injured house sparrows after a severe snow and sleet storm with high winds at Woods
individuals for a specific trait. You
hole. Of these birds 64 died, resr~ltirlgin two groups of sparrows, those that were may observe that more indfviduals
killed by the storm and the ones that survived. Bumpus, made measl~rementsof a in the population has a mean
number of randomly chosen traits such as wing length, wing span, tarsus length etc., measurement (dotted line) for the
and found that those that were killed by the storm had measurements which fell at trait.
the ends of the bell shaped curve. In other words, birds which had either mean o r b) Selection acts on individuals
with extremes of measurements
close to the rnean measurements were the ones to survive. Normalising selection (shaded areas).
generally eliminates during a catastrophe o r a stressful situation individuals whose c) As a result, the variability or
traits vary markedly from the mean values. By way of interpreting Bumpus's the population has narrowed but
observations, it could be said that the birds which were blown down easily by the the mean has remaisled the same.
winds had either their wings too long for their body weight and therefore presented
a larger surface area or they had too short wings for their body size and therefore
could not fly against strong winds. Prjor to the catastrophe the range of individual
measurements was larger as compared to the range of measurements that survived the
catastroihe. Every catastrophe may narrow down the variability of the population
thereby weeding out suboptimal genotypes.
ii) The Experiments of Dobzhansky and Spassky
Theodosius Dobzhansky and Boris Spassky demonstrated the working of normalising
selectioi~on a behavioural trait in two populations of Drosophida pseudobscura.The
 two populatior~swere subject to artificial selection. One population was selected for
positive phofobctic behaviour and the other negative phototactic behaviour Flies
were placed in a container from where they could choose to go towards light or
darkness. The photopositive flies or those that moved towards light were collected
and bred, as also tljie ones which exhibited photonegative (movement away from
light) behaviour. The brezding experiments were repeated for several generations
and for each generation the artifical selection was maintained. At the end of several
generations of artificial selection, it was observed that two distinct populations
resulted, one increasingly photopositive and the other increasingly photonegative.
More importantly, once the artificial selection was terminated natural selection
favoured individuals with neutral behaviour towards light; and both populations
reverted back to an intermediate phototactic score as shown in Fig. 11.4

Phenotype A C~ENERATIONS

1
Directional
Fig. 11.4 : Artificial selection for positive and negative phototaxis followed by normalizing natural selection
in laborntory populations of Drosophila pseudwbscum. Artificial selection for 20 generations is
successful for both negative (light triangle) and posltive (dark tria~lglcs)phototaxis. When the
Before I 1 - - afler generation 20, natural .selectionfavours a mean vnluc once again.
artificial selection is stopped
Selection
The above two example are illustrative of the fact that selection normally favoured
phenotypes which fall in the middle of the distribution, reduces variability around the
mean but does not change the mean value.

11.5.2 Action of Natural Selection in Changing Environments

1 Directional or progressive selection is the mode of selection associated with

environmental change. We said earlier that in an unchanging environment stabilising
selection removes "unusual" forms of traits or extreme measurements every
generation. But in an altered environment, the previous mean may no longer be the
optimal measurement of the trait. Under these circumstances, selection would weed
Select~on
out the individuals from the two ends of the normal curve unequally (Fig. 11,5b).
This in turn shifts the mean of the trait to a new favoured extreme (11.5c), You may
understand that directional selection is a modified form of stabilising selection in that
it does not work evenly.at both ends of the distribution curve.

Fig. 11.5 : Directional selection A revealing example of directional selection in operation is the ability of pest
a) Distribution of the population populations to develop resistance to pesticides. Man has been largely responsible for
prior to selection. changes in the environment of many organisms. In order to adapt thcmsclvcs to
b) Due to change in the changes in environment, organisms respond rapidly through directional selection.
environment,.selectbn weeds out
individuals from the two ends of
' ~ v e since
r the first report on the resistance of houseflies to DDT was made in 1947,
the curve unequally (shaded ares more and more reports of insects becoming resistant to a variety of pesticides have
indicates the individuals on whom been coming in over the past four decades. In all cases the story.is same: Initially a l
selection acts). small concentration of a new pesticide is sufficient to control thc pests. Slowly the
c) The mean value of the trait is concentration is increased even as the insects develop resistance to higher
shined to a new favoured extieme.
concentrations. Finally the pesticide becomes totally ineffective or the use of it is
economically impractical. ,
 In every generation the directoral selection eliminates from the population those me ROfg4 d Evdutionary Ch-
insects which are less and less resistant to the insecticide. Stated differently, in every
generation more and more insects that exhibit a higher resistance to the pesticide are
selected. Pesticide resistance by insects reflects the efficacy of directional selection
since the pesticides are normally synthetic substances and the insects were never
exposed to them previously in natural environments.
I I
Another interesting example of directional selection in a changing environment is
industrial melanism - a species of melanic moths escaping predation because of
altered environment. W e have a detailed discussion on this subject in Unit 12 of
Block 4. u

11.5.3 Action of Natural Selection of Heterogeneous Environments Phenotype

While describing normalising selection it was said that in uniform environments Disruptive
selection limits the variability of populations and evolves a genetic homeostasis. The Selection
type of selection process that is in progress in heterogeneous environments is known
as diversifying selection. I I
Diversifying selection is the converse of normalising selection. Let us assume that a Before
population occupying a certain environment, has two or more groups of genotypes
(AA, A a and aa) and meets two or more sub-environments or habitats. Among the
two or more genotypes, let us say that a rare genotype (aa) that is well adapted to
its habitat will be promoted by selection and the frequency of that genotype will
increase as long as the habitat is not fully occupied. But when the habitat is saturated,
no further increase in the frequency of that genotype can be observed. It is quite
likely that the excess population could spill over to another sub-environment. The
genotype may not be completely adapted to the new habitat and it is here that
diversifying selection plays a role so as to establish a population consisting different
1

k
genotypes. The phenomenon where two or more genotypes for a given trait exkt in After
a population is called genetic polymorphism. As you can see from 11.6, the different Select~on
sub-environments co'me t o be occupied by different genotypes and such an occupation
would be as complete and efficient as possible.

A.D Bradshaw and D. Jonell provided evidence that populations could become
genetically differentiated while being physically close to each other, through their
Fig. 11.6 : Disruptive selec~~on
studies on bentgrass growing on heavy metal contaminated soils. Heavy metal a) Distribution of the population
contaminants such as lead and copper are found in heaps of mine spoils and to most prior to select,on,
plants including the bentgrasses growing in the surrounding uncontaminated soils, the b) selection acts
contamination is toxic. Yet, dense growth of bentgrasses could be seen on spoil intermediate phenotypes
heaps. Essentially such plants have genes that have conferred resistance to high favours both extremes, (shaded
concentrations of lead and copper. One could observe the resistant bentgrass plants area indicates the action of
selection on individuals with
being surrounded by non-resistant varieties a few meters away in the uncontaminated htermed,ate tmils),
: soils. T h e efficiency of diversifying selection is obvious. Although cross fertilisation ,) Two distinct neighbouring
could occur between resistant and non-resistant varieties, genetic differentiation is populations each with its own
maintained because of the inability of non-resistant seedlings to grow in mean is produced.
contaiminated soil, whereas they outgrow the resistant varieties in uncontaminated
soils. Considering the fact that some of the mines are less than 400 years old it should

/ be observed that diversifying selection has produced resistant forms in a short period
oftime.

i1 SAQ3
Match the following:
I a) Selection coefficient i) measurement of reproductive efficiency
of one genotype relative to another
I1 b) Diversifying selection
genotype.
ii) Maintains genetic homeostasis.
/ C) Progressive selection iii) measure of selection pressure.
j d) Adaptive value iv) selection acting in a heterogeneous
I
environment.
i
1 e l Normalising selection v) selection that weeds away individuals
j~r
from both ends of normal curve
unequally.
i.
!
- 57
I
1.6 SUMMARY
o The principle of natural selection as put f~rwardby Damin is based on certain
facts $nd deductions. The observation that the organisms have an enormous
potential to reproduce and that in reality only a small proportion survives leads to
the deduction that there is competition among the individuals of a species for their
various as provided by the environment. This deduction when combined
with the fact that variations in heritable traits are unive1;sal in nature suggests that
in such a competition those genetic variations which are useful to the organism in
a given environment become adaptations and such adaptations improve their
reproductive efficiency. In essence the Darwinian concept of natural selection is
related to differential reproduction of the species.

o Mutations and genetic qecombinations account for the most of the variability
presented by the organisms. Both chromosomal and gene mutations occur in living
systems. Chromosomal mutations include changes in the number as well as the
structure of the chromosomes. Gene mutations, otherwise called point mutations,
bring about changes in the structure and in turn, in the function of the gene, by
the substitution or addition or deletion of a nitrogenous base. Mutation rates fol
eukaryotic genes are of the order of to per locus and prokaryotes are of
lo-' to lo-''. Sexual recombination ensures that there is a reshuffling of
chromosomes so that the genome of the zygote is to a certain extent different from
' those of the parents.

e In quantitative terms the reproductive efficiency of genotypes is measured as

fitness value. Variously termed as Darwinian fitness and adaptive value, the fitness
is an index of the reproductive efficiency of one genotype relative to alternative
genotypes within that population. The converse of fitness is selection coefficient
which is a measure of selection pressure acting on a genotype. Fitness value (w)
and selection coefficient (s) are related to each other by the expression w = (1 - s),

o Selection works on genotypes depending on the type of environments which they

occupy. If the environment is uniform and homogeneous, the type of selection that
operates is the norrnalising or stabilising kind which weeds out phenotypes with
extremesof measurements, assuming the selection pressure is very severe on such
traits, Under these circumst?nces the variability of population is reduced, but the
mean value of the trait remains the same. However, under conditions of an
environmental change, the mean value of the trait would change as the selection
acts on the two extremes of normal distribution unequally. The mean value is
shifted to a new position. The development of resistance by insects to pesticides is
an example of this type of selection known as directional selection or progressive
selection. Finally selection acting on a population distributed over a heterogene,ous
environment is known as diversifying or disrupkive selection. In heterogeneous
environment each genotype comes to occupy a sub-environment as a result of
which the normal distribution curve becomes a bimodal one with each genotype
having a separate mean value for the traits concerned.

11.7 TERMINAL QUESTIONS

1) How do you justify the statement that natural selection is synonymous with
differential reproduction?
 2) If the heritable variability does not exist in populations, could there have been Thc Process of Evolutionary Change
an evol.utionary process?
..........................................................................................................
.........................................................................................................
.........................................................................................................

3) What is meant by the term fitness value? Assuming that the selection coefficient
of a genotype is 0.35, what is the fitness value of the genotype?

.........................................................................................................
4) . What is the type of selection operating in African population in maintaining the
heterozygous genotypes HbAIHbS at a higher frequency'? (HbA - allele for
normal'haemoglobin and HbS - allele for sickle cell haenloglobin.)

.............. . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . I . . . . . . . ..........................I.................

5) A poprlatioh of isopod crustaceans Asellus nqlralicus was. found to contain two 4

variants of a gene locus coding for the enzyme amylase which breaks down starch.
Thc A, variant has optimal activity oh the starch of beech tree leaves and the Az
If v:iriant is found to have optimal activity on the starch from willow tree leaves.
The pond occupied by isopods has most of the beech trees confined to one side,
t
and the willow trees to the other side. The frequency of genotypes with beech

I1
i;
digesting amylase was found to be higher in isopods collected only at "beech
side" and the frequency of genotypes with willow digesting enzyme was higher
in collections made at the "willow side". How do you interpret the above
' situation?
 1.8 ANSWERS

i) False, (only Charles Darwin was the author)

ii) True
iii) False (They are universal in nature)
iv) False (See sub-section 3.2.2 for details)
v) True
vi) True
SAQ 2
i) Variability
ii) Mutations and genetic recombinations
iii) Inversions and translocations
iv) Polyploidy
v) Frameshift mutations
vi) Valine, glutamic acid
vii) Sickle cell trait
SAQ 3
a) iii); b) iv; c ) v; d) i; e) ii.

Terminal Questions
1) Natural selection as proposed by Darwin states that the genetic variations
produced as adaptations to changing environment are pre-requisites for the
survival of the organisms. Essentially those that survive reproduce. Among those
that reproduce, the ones that leave behind the largest number of offsprings are
regarded as the 'fittest'. Therefore the terms natural selection and differential
reproduction have similar meanings.
2) If the heritable variability does not exist in populations, then chance alone would
determine which of the organisms would survive and which would not. In the
absence of variability, essentially all organisms of a population would be uniform,
It is quite likely that any change in environment, that is adverse to the organisms,
would wipe out the entire population. Therefore, in the absence of variability,
there may not be any evolutionary change.
3) Fitness value quantifies the relative reproductive efficiency of a genotype with
reference to another genotype. If s, the selection coefficient of a genotype is 0.35,
then w , the fitness value of the genotype is 1-0.35 = 0.65.
4) In African populations, selection maintains the hetrozygotes HbA/HbS at high
frequencies. The two homozygotes are not favoured for the following reasons.
The HbS/HbS genotype causes the sickle all disease and the persons homozygous
for HbS die early because of the lysis of erythrocytes. (Refer section 11.3.2).
HbAjHbA homozygotes are susceptible to falciparum malaria. In hetrozygotes
where an HbS allele is present, there is the sickling of cells to a certain extent,
but the genotypes exhibit resistance to malaria. Since both the homozygotes are
selected against and only heterozygotes favoured, it is the normalising selection
that is operating in African population.
5) The example illustrates the action of diversifying selection. In the pond there arc
two isopod populations differing from one another in the type of amylase they
possess. The isopods come to occupy heterogeneous environments, one
population confined to 'beach side' and the other the 'willow side'. Natural
selection has separated the two genotypes (they are two genotypes because of the
differences in the amylase activity of the two populations) but retained them in
the same neighbouring environments.

GLOSSARY ,

Acridines : a group of aromatic dyes whose dimensions are same as a nitrogenous ,

bnse pair. They wedge between purines and pyrimidines of intact DNA inducing ,
distortions in the helix causing additionsor deletions. Acridine orange is a well knowd
60 mutagenic agent. j
I
I
I
 Adaptive radiation : the tendency of successful species to diversify into all types of The Process of Evolutionary Change

ecological niches.
Biogeography : the study o f the distribution of organisms over the earth and of the
principles that govern their distribution.
Carrying capacity : the population density that can be sustained by limiting sources.
Chondrodystrophy : a hereditary disease causing the deformation of skeletal system.
Endemic : refers to organisms which are narrowly restricted in their distribution.
Epoch : a unit of geological time; it is the principal subdivision of pc~iods.
Fossil : petrified remains of the organisins of the past.
Gene pool : the total of all genes possessed by reproductive members of a population.
Geology : study of the science of earth.
Huntington's chorea : a rare hereditary disease affecting brain resulting in choreiform
(dance like) movement, intellectual deterioration and psychosis.
Isotopes : elements having the same atomic number but different atomic weight.
Monozygotic twins : twins produced from a single fertilised egg. The first division of
the zygote produces two cells, each of which develops into an embryo; also known
as identical twins.
Mutation : an inheritable change in gene.
Palaeontology : the study of the life of the past through fossil records.
Pangenesis : a theory of inheritance proposed by Darwin according to which all organs
in the body produce pangenes, that is minute particles that are carried away by the
blood stream and segregated out into gametes.
Pentadactyly : refers to the presence of five digits in limbs, very characteristic of
higher vertebrates.
Positive phototactism : ability to respond to light or move in the direct~onof light.
This is in contrast t o negative phototropism in which organisms tend to avoid light
and move towards darkness,
PTC taster : phenotypes who can taste a solution of phenyl thiocarbamide bitter. The
ability to taste the solution is controlled by a single dominant gene. PTC non-tasters
do not have any taste to the chemical.
Radioactive dating : determining the age of geological deposits and fossils based on
the rate of decay of radioactive elements.
Recombination : a phenomenon occurring at the time of meiosis leading to the
shuffling of chromosomes and as a result the genes as well.
Stratigraphy : the study of layers of rock with reference to relative age of forms
contained in them.
' Tautomeric shifts : a reversible change in a molecule brought about by a shift in the
location of a hydrogen atom. In nucleic acids tautomeric shifts in the bases of
nucleotides can cause changes in other bases during replication and are a source of
mutation.
) Ungulates : hoofed mammals.

/1 Vicariance : A pattern of distribution in which related populations replace each other

in widely separated areas.

i FURTHER READINGS
I) Dodson, E.O. (1985) Evolution: Process and Product. Wadsworth Publishing
b Company, California, USA.
2) Lull, R.S.(1984) Organic Evolution(Revised edition). Seema Publications, Delhi.
: 3) Moody, P.A. (1978) Intloduction to Evolution (Third edition), i<alyani
Publishers, Delhi.
4) Stebbins G.L. Jr. (1968) Variation and Evolution in Plants. Oxford and IBH
Publishing Co. Calcutta.
i
Dear Student,

.While studying these units you may have found cer,tain portions of the text difficult to
comprehend. We wish to know your difficulties and suggestions in order to improve 'the
course. Therefore, we request you to fill and send us the following questionnaire which
pertains t o this block.

QUESTIONNAIRE

1) How may hours did you need for studying the units?

Unit Number
No. of hour*

2) How many hours (approximately) did you take to do the assignments pertaining to
this block?

Assignment Number
No. of hours.

3) In the following table we have listed 4 kinds of difficulties that we thought you might
have come across. Kindly tick (V) the type of difficulty and give the relevant page
number in the appropriate columns.

Page Number Types of difficulties

Presentation Language Diagram Terms are
is not clear is difficult is not clear not explained

1
4) It is possible that you could not attempt some SAQs and TQs.
In the following table are listed the possible difficulties. Kindly tick (V) the type of
difficulty and tlie relevant unit and question numbers in the appropriate columns.

Type of difficulty
-
Unit 'SAQ . TQ Not clearly Cannot answer Answer given Answer
No. No. No. ' posed on basis of (at end of given is
information Unit) not
given not clear sufficient

sj Were all the difficult terms included in the glossary. If not, please list in the space
given below.
 Any other suggestion(s)

Affix
Postage
Stamp

To
The Course Coordinator
(LSE-07, Taxonomy and Evolution)
School of Sciences
Indira Gandhi National Open University
Maidan Garhi
New Delhi-110 068
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