MZO-004

SYSTEMATICS,
BIODIVERSITY
AND EVOLUTION

VOLUME 1 SYSTEMATICS
(Blocks 1 and 2)
 MZO-004
SYSTEMATICS, BIODIVERSITY
Indira Gandhi AND EVOLUTION
National Open University
School of Sciences

VOL

1
SYSTEMATICS
BLOCK 1
EVOLUTIONARY BIOLOGY-I 5
BLOCK 2
EVOLUTIONARY BIOLOGY-II 45
 MZO-004
SYSTEMATICS, BIODIVERSITY
Indira Gandhi AND EVOLUTION
National Open University
School of Sciences

Block

1
EVOLUTIONARY BIOLOGY-I
UNIT 1
Concept of Organic Evolution 9
UNIT 2
The RNA World and Origin of Life 27
COURSE NAME: SYSTEMATICS, BIODIVERSITY AND EVOLUTION COURSE CODE: MZO-004

Course Design Committee

Prof. Sujatha Varma Prof. Bharati Chauhan Prof. Amrita Nigam (Retd.)
Former Director, School of Department of Zoology School of Sciences, IGNOU
Sciences, IGNOU, Maidan Garhi University of Rajasthan Maidan Garhi, New Delhi-110068
New Delhi-110068 Jaipur-302004
Prof. Neera Kapoor
Prof. Neeta Sehgal Prof. Devinder Kaur Kochar School of Sciences, IGNOU
Department of Zoology Department of Zoology, Punjab Maidan Garhi, New Delhi-110068
University of Delhi, Delhi-110007 Agricultural University
Ludhiana-141004 Dr. Siya Ram
Prof. Sarita Kumar School of Sciences, IGNOU
Department of Zoology, Acharya Prof. Varsha Wankhade Maidan Garhi, New Delhi-110068
Narender Dev College, University Department of Zoology
of Delhi, Delhi-110007 Savitribai Phule University Dr. Ravi Rajwanshi
Pune-411007 School of Sciences, IGNOU
Prof. S. Dinakaran Prof. Sarita Sachdeva Maidan Garhi, New Delhi-110068
Department of Zoology Department of Biotechnology
Madurai Kamaraj University Manav Rachna International
Tamil Nadu-625021 University, Sector-43, Faridabad
Haryana-121004

Block Preparation Team

Dr. Anjali S. Nawani School of Sciences
Department of Zoology,
Dr. Ravi Rajwanshi
Sri Venkateswara College,
School of Sciences, IGNOU
South Campus,University of Delhi,
Maidan Garhi, New Delhi-110068
Delhi 110021 (Unit-1 and 2)
(Units-1 and 2)

Programme Coordinators : Prof. Neera Kapoor and Dr. Siya Ram

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Course Editor : Dr. Tapati Das,
Associate Professor,
Department of Ecology and Environmental Sciences,
Assam University, Silchar 788011, Assam

Production : Mr. Tilak Raj

AR, MPDD, IGNOU
Acknowledgement:
• Dr. Ravi Rajwanshi and Mr. Ajit Kumar - Suggestions for figures and cover design.

• Mr. Manoj Kumar, Assistant for word processing and CRC preparation.
August, 2024
Indira Gandhi National Open University, 2024
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Unit 1 Introduction to Evolutionary Biology

UNIT 1
INTRODUCTION TO
EVOLUTIONARY BIOLOGY

Structure
1.1 Introduction 1.4 Charles Darwin and his
Theory of Evolution
Objectives
1.5 Post Darwinian Era
1.2 What is Evolution?
Neo-Lamarckism
1.3. Pre Darwinian Era
Theory of Orthogenesis
Plato’s Philosophy of
Essentialism Mutanist Theory

Aristotle’s Idea of Species Evolutionary synthesis

with Fixed Properties
1.6 Fundamental principles of
Theory of Special Creation Evolution
Great Chain of Being or 1.7 Philosophical Issues
Scala Nature Related to Darwin’s
Evolutionary Theory
Classification in His Systema
Naturae 1.8 Summary
Principle of Uniformitarianism 1.9 Terminal Questions
Inheritance of Acquired 1.10 Answers
Characteristics

2.1 INTRODUCTION
When we look around ourselves we see a variety of organisms/ species, all of
which are different from one another. You must be amazed to see how do
peacocks have such beautiful feathers? Why do whales have lungs, and why
do snakes lack legs? Why does one species of ant have a single
chromosome, while some butterflies have more than 200? How this variation
in living forms can be answered?

In one of the most breathtaking ideas in the history of science, Charles Darwin
proposed that "all the organic beings which have ever lived on this earth have
descended from one primordial form." From this idea, it follows that every
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Block 1 Evolutionary Biology-I

characteristic of every species- the feathers of a peacock, the number and

sequence of its genes, the catalytic abilities of its enzymes, the
structure of its cells and organs, its physiological tolerances and nutritional
requirements, its life span and reproductive system, its capacity for behaviour
is the outcome of an evolutionary history. The evolutionary perspective
illuminates every subject in biology, from molecular biology to ecology. Indeed,
evolution is the unifying theory of biology. The geneticist Theodosius
Dobzhansky appropriately quoted that "Nothing in biology makes sense,
except in the light of evolution”.

This is the first unit of the course and we are going to discuss about the basic
concept of evolution, its emergence and acceptance in society.

Objectives
Upon completion of this unit, you would be able to:
 define of evolution;

 comprehend the originator of the concept and its societal acceptance;

and

 Describe the emergence of the concept of evolution and its role in

explaining variations among living organisms.

1.3 WHAT IS EVOLUTION?

The word evolution comes from the Latin word ‘evolvere’, which means to
unfold or to reveal or manifest hidden potentialities. Evolution is the change in
the properties of groups of organisms over the course of generations. The
development, or ontogeny of an individual organism is not considered
evolution while groups of organisms, which we call population, undergo
evolution or modification. The changes in populations that are considered
evolutionary are those that are passed via the genetic material from one
generation to the next.

A current example of evolution is the trend of antibiotic resistance in microbial

population. Before the 1940s, people suffering from some bacterial infections
like tuberculosis and pneumonia had little hope of getting cured. However by
1965, the discovery of antibiotic drugs and subsequent advances in their
synthesis led to the conquest of such bacterial diseases. But at present most
bacteria are becoming resistant to antibiotics like penicillin, ampicillin, that
were supposed to have beat them out.

Staphylococcus aureus, for example, a bacterium that causes many infections

in surgical patients, is now almost universally resistant to penicillin, ampicillin,
and related drugs. Subsequently, Methicillin was developed as an alternative
which worked for a few years. However, in due course of time S. aureus
became resistant to methicillin and then to other antibiotics like
cephalosporins, carbapenelns, erythromycin, tetracycline, streptomycin,
sulfonamides, and fluoroquinolones. This was followed by discovery of another

10
Unit 1 Introduction to Evolutionary Biology

new drug, varicomycin which seemed to have solved the problem, but it too is
becoming less effective.

As the use of antibiotics increases, so does the incidence of bacteria

becoming resistant to those antibiotics. Why is this happening? Do the
excessive use of drugs cause drug-resistance in the bacterial genes? How it
works? Does the resistance spread from one bacteria to another? Can the
resistance of the bacteria be prevented by using lower or higher doses of
drugs or by combinations of different drugs?

The principles and methods of evolutionary biology have provided some

answers to these questions, and to many others that affect society. Moreover,
there are many other questions that can be answered by applying evolutionary
principles, some of them are mentioned below:

• the transfer of HIV to humans from chimpanzees and old world

monkeys.

• the development of insecticide resistance in insects that spread disease

and ruin crops.

• the methods for genetically transferring disease and insect resistance

from wild plants to crop plants, as well as non-chemical pest control
methods.

• designing new drugs and other usefull products.

• In computer science and artificial intelligence, ‘evolutionary computation’

uses principles taken directly from evolutionary theory to solve
mathemalically intractable practical problems, such as constructing
complex timetables or processing radar data.

The importance of evolutionary biology goes far beyond its practical uses. The
way we think about ourselves can be profoundly shaped by an evolutionary
framework. How do we account for human variation-the fact that almost
everyone is genetically and phenotypically unique? What about apparently
useless or even potentially harmful characteristics such as our wisdom teeth
and appendix? Why does noncoding apparently useless DNA account for
more than 98 percent of the human genome? Why do we age, undergo
senescence, and eventually die? Why are medical researchers able to use
monkeys, mice and even fruit flies and yeasts as models for processes in the
human body? Such questions and their answers lie in the realm of evo-
lutionary biology.

SAQ 1
a) Fill in the blanks:

i) The word evolution comes from the Latin evolvere, means……….

ii) In biology, evolution is defined as……………………

iii) Evolutionary principles are applied to a……………not to an

individual organism.
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Block 1 Evolutionary Biology-I

iv) The changes in populations that are considered evolutionary are

those that are passed via …………..from one generation to the
next.

b) Give a current example of evolution in nature.

1.3 PRE DARWINIAN ERA

The Pre-Darwinian era, spanning centuries before Charles Darwin's
groundbreaking work on evolution, represents a period of diverse scientific
ideas and philosophical contemplation about the origins and diversity of life on
Earth. Prior to Darwin's publication of "On the Origin of Species" in 1859,
prevalent beliefs about the natural world were deeply influenced by religious
doctrines, ancient texts, and limited empirical observations. Pre-Darwinian era
was marked by a rich tapestry of scientific inquiry, religious interpretation, and
philosophical speculation regarding the origins and diversity of life which are
briefly described as follows:

Plato’s philosophy of essentialism: The guiding philosophical idea about

humans in particular and biology in general into the 19th Century was Plato’s
concept of ‘Essentialism’. Plato believed there existed ideal forms of
everything, including a human. Variation is attributed to
the imperfections manifested in individuals (defects of the underlying perfect
essence/form).

Aristotle’s idea of species with fixed properties: Plato's philosophy of

Plato’s essentialism
essentialism became incorporated into Western philosophy largely through
worked well in
physics and Arlstotle (Aristotle was the student of Plato), who developed Plato's concept of
chemistry, and it immutable essences into the notion that species have fixed properties.
helped scientists
discover some of the Theory of special creation: Later, Christians interpreted the biblical account
mathematical of ‘Genesis’ and concluded that each species had been created individualy by
formulas (the perfect God in the same form it has today.
forms) that underlie
all that is. So there Great Chain of Being or scala nature: Christian thought elaborated on
was good reason for Platonic and Aristotelian philosophy, arguing that since existence is good and
biology, trying to be a God's benevolence is complete, He must have bestowed existence on every
hard science, to creature, each with a distinct essence, of which he could conceive. Because
embrace essentialism
order is superior to disorder God's creation must follow a plan: specifically, a
as well.
gradation from inanimate objects and barely animate forms of life, through
plants and invertebrates, up through ever "higher" forms of life. Humankind,
which is both physical and spiritual in nature, fanned the link between animals
and angels. This "Great Chain of Being," or scala nature (the scale, or ladder,
of nature), must be permanent and unchanging, since change would imply that
there had been imperfection in the original creation.

Classification in Carolus Linnaeus Systema Naturae: Carolus Linnaeus

(1707-1778), who established the framework of modern classification in his
Systema Naturae (1735), undertaken in the hope of discovering the pattern of
the creation won worldwide fame for his exhaustive classification of plants and
12
Unit 1 Introduction to Evolutionary Biology

animals,. Linnaeus classified "related" species into genera, "related" genera

into orders, and so on. To him, "relatedness" meant propinquity in the
Creator's design.

Principle of Uniformitarianism: James Hutton, an 18th-century Scottish

geologist, is often credited with formulating this principle, though its ideas can
be traced back to earlier thinkers. Hutton emphasized the gradual
accumulation of geological changes over immense periods of time. The
Principle of Uniformitarianism is a foundational concept in geology and earth
sciences, proposing that the same natural laws and processes that operate in
the present have also operated in the past, and will continue to operate in the
future.

Inheritance of Acquired Characteristics: The most significant pre-Darwinian

evolutionary hypothesis, representing the culmination of eighteenth-century
evolutionary thought, was proposed by the Chevalier de Lamarck in his
Philosophie Zoologique (1809). Lamarck proposed that each species
originated individually by spontaneous generation from non-living matter
starting at the bottom of the chain of being. He mentioned that a "nervous
fluid" acts within each species, causing it to progress up to the chain. As
species vary in age and have distinct origins, we currently see a hierarchy of
species.

Lamarck argued that species differ from one another because they have
different needs, and so use certain of their organs and appendages more than
others. The more strongly exercised organs attract more of the "nervous fluid,"
which enlarges them, just as muscles become strengthened by work.
Lamarck, like most people at that time, believed that such alterations, acquired
during an individual's lifetime, are inherited - a principle called ‘inheritance of
acquired characteristics’. In the most famous example of Lamarck's theory,
giraffes originally had short necks, but stretched their necks to reach foliage
above them. Hence their necks were lengthened; longer necks were inherited,
and over the course of generations, their necks got longer and longer. This
could happen to any and all giraffes, so the entire species could have acquired
longer necks because it was composed of individual organisms that changed
during their lifetimes.

Lamarck's ideas had little impact during his lifetime, partly because they were
criticized by zoologists at that time, and partly because after the French
Revolution, ideas originating from France were suspected in most other
countries. Though Lamarck's ideas of how evolution works were wrong, he
deserves credit for being the first to advance a coherent theory of evolution.

SAQ 2
a) What is philosophy of essentialism? Who proposed it?

b) What is theory of special creation?

c) What does Scala nature represents?

d) Who gave the first systemic classification of animals and plants?

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Block 1 Evolutionary Biology-I

e) What is Lamark’s theory of “Inheritance of acquired chracterstics“?

1.4 CHARLES DARWIN AND HIS THEORY OF

EVOLUTION
Charles Robert Darwin (February 12, 1809-April 19, 1882) was the son of an
English physician. He briefly studied medicine at Edinburgh. After that he
turned to studying for a career in the clergy at Cambridge University. He
apparently believed in the literal truth of the Bible as a young man. He was
passionately interested in natural history and became a companion of the
natural scientists. His life was changed forever, when he was invited to serve
as a naturalist on the H.M.S. Beagle, a ship the British navy was sending to
map the waters of South America in 1831.

The Beagle's voyage lasted from December 27, 1831 to October 2, 1836. The
ship spent several years traveling along the coast of South America, where
Danwin observed the natural history of the Brazilian rain forest and the
Argentine pampas, and stopped in the Galapagos Islands (on the Ecuator off
the coast of Ecuador). In the course of the voyage, Darwin became an
accomplished naturalist, collected specimens, made innumerable geological
and biological observations, and conceived a new (and correct) theory of the
formation of coral atolls. Soon after his return, the ornithologist John Gould
pointed out that Darwin's specimens of mockingbirds from the Galapagos
Islands were so different from one island to another that they represented
different species. Darwin then recalled that the giant tortoises, too, differed
from one island to the next. These facts, and the similarities between fossil
and living mammals that he had found in South America, triggered his
conviction that different species had evolved from common ancestors.

Darwin's comfortable finances enabled him to devote the rest of his life
exclusively to his biological work (although he was chronically ill for most of his
life after the voyage). He set about a massive evidence of evolution and tried
to conceive of its causes. On September 28, 1838, he read an essay by the
economist Thomas Malthus, who argued that the rate of human population
growth is greater than the rate of increase in the food supply, so that
unchecked growth must lead to famine. This was the inspiration for Darwin's
great idea, one of the most important ideas in the history of thought: ‘Natural
selection’. Darwin wrote in his autobiography that "being well prepared to
appreciate the struggle for existence which everywhere goes on from long-
continued observation of the habits of animals and plants, it at once struck me
that under these circumstances favourable variations would tend to be
preserved and unfavourable ones to be destroyed." In other words, if
individuals of a species with superior features survived and reproduced more
successfully than individuals with inferior features, and if these differences
were inherited, the average character of the species would be altered.

Mindful of how controversial the subject would be, Darwin then spent twenty
years amassing evidence about evolution and pursuing other researches
14 before publishing his ideas. He wrote a private essay in 1844, and in 1856 he
Unit 1 Introduction to Evolutionary Biology

finally began writing a book he intended to call Natural Selection. He never

completed it, for in june 1858 he received a manuscript from a young
naturalist, AIfred Russel Wallace (1823-1913). Wallace, who was collecting
specimens in the Malay Archipelago, had independently conceived of natural
selection. Darwin had extracts from his 1844 essay presented orally, along
with Wallace's manuscript, at a meeting of the major scientific society in
London, and set about writing an "abstract" of title book he had intended.
Darwin became well-known and controversial almost immediately after the
publication of his 490-page "abstract," which was titled "On the Origin of
Species by Means of Natural Selection," or "The Preservation of Favoured
Races in the Struggle for Life." It was published on November 24, 1859.

For the rest of his life, Darwin continued to read and correspond on an
immense range of subjects, to revise The Origin of Species (it had six
editions), to perform experiments of all sorts (especially on plants), and to
publish many more articles and books, of which The Descent of Man is the
most renowned.

The Origin of Species has two major theses. The first is Darwin's theory of
“descent with modification”. It holds that all species, living and extinct, have
descended, without interruption, from one or a few original forms of life.
Species that diverged from a common ancestor were at first very similar, but
accumulated differences over great spans of time, so that some are now
radically different from one another. Darwin's conception of the course of
evolution is profoundly different from Lamarck's, in which the concept of
common ancestry plays almost no role.

The second theme of The Origin of Species is Darwin's theory of the causal
agents of evolutionary change. This was his “theory of natural selection”: "if
variations useful to any organic being ever occur, assuredly individuals thus
characterised will have the best chance of being preserved in the struggle for
Iife; and from the strong principle of inheritance, these will tend to produce
offspring similarly characterised. This principle of preservation, or the survival
of the fittest is called as natural selection."

Darwin's theory of evolution actually includes five theories:

1. Evolution as such is the simple proposition that states that

characteristics of lineages of organisms change over time. This idea was
not original with Darwin, but it was Darwin who so convincingly
marshaled the evidence for evolution that most biologists soon accepted
that it has indeed occurred.

2. Common descent is a radically different view of evolution than the

scheme Lamarck had proposed. Darwin was the first to argue that
species had diverged from common ancestors and that all of life could
be portrayed as one great family tree.

3. Gradualism in Darwin's proposition states that the differences between

even radically different organisms have evolved incrementally, by small
steps through intermediate forms. The alternative hypothesis is that
large differences evolve by leaps, or saltations, without intermediates.
15
Block 1 Evolutionary Biology-I

4. Population change in Darwin's thesis states that evolution occurs by

changes in the proportions of individuals within a population that have
different inherited characteristics. This concept was a completely original
idea that contrasts both with the sudden origin of new species by
saltation and with Lamarck's account of evolutionary change by
transformation of individuals.

5. Natural selection was Darwin's brilliant hypothesis, independently

conceived by Wallace, that changes in the proportions of different types
of individuals are caused by differences in their ability to survive and
reproduce and that such changes result in the evolution of adaptations,
features that appear "designed" to fit organisms to their environment.
The concept of natural selection revolutionized not only biology, but
Western thought as a whole.

Darwin proposed that the various descendants of a common ancestor evolve

different features because they are adaptive under different "conditions of life"-
different habitats or habits. Moreover, the pressure of competition favors the
use of different foods or habitats by different species. He believed that no
matter how extensively a species has diverged from its ancestors, new
hereditary variations continue to arise, so that given enough time, there is no
evident limit to the amount of divergence that can occur.

From where do these hereditary variations come? This was the great gap in
Darwin's theory, which he never could fill it. The problem was serious,
because according to the prevailing belief in Blending Inheritance, variation
should decrease, not increase. Because offspring are often intermediate
between their parents in features such as color or size, it was widely believed
that characteristics are inherited like fluids, such as different colors of paint.
Blending white and black paints produces gray, but mixing two gray paints
doesn't yield black or white: variation decreases. Darwin never knew that
Gregor Mendel had infact solved the problem in a paper published in 1865.
Mendel's theory of Particulate Inheritance proposed that inheritance is based
not on blending fluids, but on particles that pass unaltered from generation to
generation so that variation can persist. The concept of "mutation" in such
particles (later called genes) developed only after 1900 and was clarified
considerably later.

SAQ 3
a) Who was Charles Darwin?

b) Charles Darwin was impressed by the essay on population growth

written by which economist?

c) What is the meaning of “evolution by common descent”?

d) According to Charles Darwin what is the cause of evolution?

1.5 POST DARWINIAN ERA

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Unit 1 Introduction to Evolutionary Biology

Although The Origin of Species raised enormous controversy, by the 1870s

most scientists accepted the historical reality of evolution by common descent.
Thereafter, in the late nineteenth and early twentieth centuries, evidence from
paleontology, comparative morphology, and comparative embryology studies
were found, which adds lots of information on evolution in the fossil record and
on relationships among organisms. But this consensus did not extend to
Darwin's theory of the cause of evolution, natural selection. For about 60 years
after the publication of The Origin of Species, many scientists rejected natural
selection as the cause of evolution, and numerous theories were proposed
instead. These theories included neo-Lamarckian, orthogenetic, and
mutationist theories (Bowler 1989).

Neo-Lamarckism includes several theories based on the old idea of

inheritance of modified acquired characters during an organism's lifetime.
Such modifications might have been due to the direct effect of the environment
on developlnent (as in plants that develop thicker leaves if grown in a hot, dry
environnlent). In a famous experiment, August Weismann cut off the tails of
mice for many generations and showed that this had no effect on the tail
length of their descendants. Subsequently, extensive research provided no
evidence that specific hereditary changes can be induced by environmental
conditions under which they would be advantageous.

Theories of Orthogenesis, or "straight-line evolution," held that the variation

that arises is directed toward fixed goals, so that a species evolves in a
predetermined direction without the aid of natural selection. Some
paleontologists held that such trends need not be adaptive and could even
drive species towards extinction. None of the proponents of orthogenesis ever
proposed a mechanism for it.

Mutanist theories were advanced by some geneticists who observed that

discretely different new phenotypes can arise by a process of mutation. They
supposed that such mutant forms constituted new species, and thus believed
that natural selection was not necessary to account for the origin of species.
Mutationist ideas were advanced by Hugo de Vries, one of the biologists who
"discovered" Mendel's neglected paper in 1900, and by Thomas Hunt
Morgan, the founder of Drosophila genetics. The last influential mutationist
was Richard Goldschmidt (1940), an accomplished geneticist who
nevertheless erroneously argued that evolutionary change within species is
entirely different in kind from the origin of new species and higher taxa.
According to him these originated by sudden, drastic changes that reorganize
the whole genome. Although most such reorganisations would be deleterious,
a few "hopeful monsters" would be the progenitors of new groups.

The Evolutionary Synthesis

These anti-Darwinian ideas were refuted in the 1930s and 1940s by the
evolutionary synthesis or modern synthesis, forged from the contributions
of geneticists, systematists, and paleontologists who reconciled Darwin's
theory with the facts of genetics. A list of scientists and their contributions in
the field of studies related to evolution is presented below:

Scientists Field of study Contributions

17
Block 1 Evolutionary Biology-I
Ronald A. Fisher, John B. Developed a Showed that mutation and natural
S. Haldane, Sewall mathematical selection together cause adaptive
Wright theory of evolution: mutation is not an
population alternative to natural selection! but
genetics is rather its raw material.
Sergei Chetverikov Population Study of genetic variation and
Genetics change in natural populations
Theodosius Dobzhansky Population Wrote an influential book titled
Genetics “Genetics and The Origin of
Species” in 1937. He conveyed the
ideas of the population geneticists
to other biologists, thus influencing
their appreciation of the genetic
basis of evolution.
Ernst Mayr Zoologist Wrote book titled “Systematics ond
The Origin of Species” in (1942)
Bernhard Rensch Ornithologist Evolution Above the Species Level
(1959)
G. Ledyard Stebbins Botanist Variation and Evolulian in Plants
(1950)
George Gaylord Simpson Paleontologist Tempo and Mode in Evolution
(1944) and The Major Features of
Evolution (1953)

These authors argued persuasively that mutation, recombination, natural

selection, and other processes operating within species (which Dobzhansky
termed microevolution) account for the origin of new species and for the
major, long-term features of evolution (termed macroevolution).

SAQ 4
State True or False:

a) During Post Darwinian Era by 1870’s most scientists accepted the

historical reality of evolution by common descent.

b) According to Darwin's theory, the cause of evolution is natural selection.

c) For many years, many scientists rejected natural selection as the cause
of evolution.

d) Neo-Lamarckism includes theories related to inheritance of acquired

chrachterstics as the process of evolution.

e) Mutanist theories proposed discretely different new phenotypes can

arise by a process of mutation and it may be the leading cause of
evolution.

f) Theories of Orthogenesis, or "straight-line evolution," held that the

variation that arises is directed toward fixed goals, so that a species
evolves in a predetermined direction without the aid of natural selection.

18
Unit 1 Introduction to Evolutionary Biology

g) The evolutionary synthesis or modern synthesis, rejects Darwin's

theory of evolution (and its cause as Natural Selection) using the facts of
genetics.

1.6 FUNDAMENTAL PRINCIPLES OF EVOLUTION

The principle claims of the evolutionary synthesis are the foundations of
modern evolutionary biology. Although some of these principles have been
extended, clarified, or modified since the 1945, most evolutionary biologists
today accept them as fundamentally valid. These are the fundamental
principles of evolution which are discussed at length throughout this course.
The table below shows the basic principles of evolution and the related
explanation(s).

S.No. Principles of Evolution Explanation

1. The phenotype (observed Phenotypic differences among individual

characteristic) is different from organisms may be due partly to genetic
the genotype (the set of genes differences and partly to direct effects of the
in an individual's DNA). environment.
2. The acquired characteristics Environmental effects on an individual's
are not inherited. phenotype do not affect the genes passed
on to its offspring.
3. Hereditary variations (whether Genes retain their identity as they pass
discrete or continuous) are through the generations; they do not blend
based on genes. with other genes. This is true of both
discretely varying traits (e.g., brown vs. blue
eyes) and continuously varying traits (e.g.,
body size, intensity of pigmentation).
Genetic variation in continuously varying
traits is based on several or many discrete,
particulate genes, each of which affects the
trait slightly ("polygenic inheritance").
4. Genes mutate to equally Gene mutation takes place usually at a fairly
stable altemative forms, low rate. The phenotypic effect of such
known as alleles. mutations can range from undetectable
change to very great change. The variation
that arises by mutation is amplified by
recombination among alleles at different
loci.
5. Evolutionary change is a Evolution entails, in its most basic form, a
populational process. change in the relative abundances
(proportions or frequencies) of individual
organisms with different genotypes (hence,
often, with different phenotypes) within a
population. One genotype may gradually
replace other genotypes over the course of
generations. Replacement may occur within
only certain populations, or in all the
populations that make up a species.
6. The rate of mutation is too low Instead, the change in genotype proportions
to shift a population from one within a population can occur by either of
19
Block 1 Evolutionary Biology-I
genotype to another. two principal processes: random fluctuations
in proportions (genetic drift), or non-
random changes due to the superior survival
and/or reproduction of some genotypes
compared with others (i.e., natural
selection). Natural selection and random
genetic drift can operate simultaneously.
7. Natural selection can account Even a slight intensity of natural selection
for both slight and great can (under certain circumstances) bring
differences among species, as about substantial evolutionary change in a
well as for the earliest stages realistic amount of time.
of evolution of new traits.
Natural selection can alter populations
beyond the original range of variation by
increasing the frequency of alleles that, by
recombination with other genes affect the
same trait leading to generation of new
phenotypes.
Natural populations are genetically variable
and so can often evolve rapidly when
environmental conditions change.
8. The differences between The differences between different species,
species evolve in small steps. and between different populations of the
same species, are often based on
differences at several genes, many of which
have a small phenotypic effect.
Among living organisms, there are many
gradations in phenotypic characteristics
among species assigned to the same
genus, to different genera, and to different
families or other higher taxa. Such
observations provide evidence that higher
taxa arise by the prolonged, sequential
accumulation of small differences, rather
than by the sudden mutational origin of
drastically new "types."
9. Adaptations are traits that Populations of a species in different
have been shaped by natural geographic regions differ in characteristics
selection. that have a genetic basis.
Differences among geographic populations
of a species are often adaptive, and thus are
the consequence of natural selection.
10. A species is a group of Phenotypically different genotypes are often
interbreeding or potentially found in a single interbreeding population.
interbreeding individuals that Species are not defined simply by
do not exchange genes with phenotypic differences. Rather, different
other such groups. species represent distinct "gene pools".
Speciation is the origin of two or more
species from a single common ancestor.
Speciation usually occurs by the genetic
differentiation of geographically segregated
populations. Because of the geographic
segregation, interbreeding does not prevent
incipient genetic differences from
20
Unit 1 Introduction to Evolutionary Biology
developing.
11. The evolution of large The fossil record includes examples of
differences, which leads to gradations from apparently ancestral
macroevolution, proceeds organisims to quite different descendants.
incrementally. However, the complete fossil record for
quite different kinds of organisms are not
available at many times. But they support
the hypothesis that the evolution of large
differences proceeds incrementally. This
also substantiate the fact that the principles
that explain the evolution of populations and
species may be extrapolated to the
evolution of higher taxa.

Evolutionary Biology since the synthesis

Since the evolutionary synthesis involves a great deal of research has

elaborated and tested its basic principles. Since the 1950s, the study of
evolution has undergone a virtual revolution due to the advancements in
genetics and molecular biology, which have also opened up completely new
research areas like molecular evolution. This revolution has been intensifying
ever since. Molecular biology has provided tools for studying a vast number of
evolutionary topics, such as mutation genetic variation, species differences,
development, and the phylogenetic history of life.

Since the mid-1960s, evolutionary theory has expanded into areas such as
ecology, animal behavior, and reproductive biology, and detailed theories have
been developed to explain the evolution of particular kinds of characteristics
such as life span, ecological distribution, and social behavior. The study of
macroevolution has been renewed by provocative interpretations of the fossil
record and by new methods for studying phylogenetic relationships. As
molecular methods have become more sophisticated and available, virtually
new fields of evolutionary study have developed. Among these fields is
Molecular Evolution (analyses of the processes and history of change in
genes), in which the Neutral Theory of Molecular Evolution has been
particularly important. This hypothesis, developed especially by Motoo Kimura
(1924 -1994), holds that most of the evolution of DNA sequences occurs by
genetic drift rather than by natural selection. Evolutionary Developmental
Biology is an exciting field devoted to understanding of how developmental
processes have both evolve and constrain evolution. It is closely tied to
developmental biology, one of the most rapidly moving fields of biology today.

Evolutionary Genomics, which is an emerging field focuses on the mutation

and evolution of multiple genes or entire genomes. The progress in these
areas is supported by vigorous research, new discoveries, and innovative
ideas about long-standing topics in evolutionary biology, such as the evolution
of adaptations and the emergence of new species. This is a thrilling time to
study evolution or to be an evolutionary biologist.

21
Block 1 Evolutionary Biology-I

SAQ 5
Fill in the blanks:

a) The phenotype (observed characteristic) is …………….. from the

genotype (the set of genes in an individual's DNA).

b) The acquired charachterstics are ……………inherited.

c) Hereditary variations (wheather discrete or continuous) are based on

…………..….

d) Genes mutate to equally stable altemative forms, known as ………….

e) Evolutionary change is a …………….. process.

f) Adaptations are traits that have been shaped by………………

1.7 PHILOSOPHICAL ISSUES RELATED TO

DARWIN’S EVOLUTIONARY THEORY
Many articles have been written about the philosophical and social
implications of evolution. Few major objections are mentioned here:

1. Rejection of Essentialism: Darwin argued that, given enough time, a

species' entire range of characteristics might undergo drastic alteration.
As a result, he substituted variation for the essentialism theory that
Western philosophy had inherited from Plato and Aristotle.

2. Question mark on God’s perfect creation which is static: Darwin

also helped to replace a static conception of the world - one virtually
identical to the creators perfect creation with a world of ceaseless
change. It was Darwin who extended to living things, including the
human species, the principle that change, not stasis, is the natural order.

3. Lack of purpose in natural selection: Above all, Darwin's theory of

random, purposeless variation acted on by blind, purposeless natural
selection provided a revolutionary new kind of answer to almost all
questions that begin with "Why?". Prior to Darwin, purpose was used by
philosophers and people in general to respond to "Why?" queries.

4. Since only an intelligent mind, with the capacity for forethought, can
have purpose, questions such as why do plants have flowers?" or "Why
are there apple trees?"--or diseases, or earthquakes were answered by
imagining the possible purpose that God could have had in creating
them. The kind of explanation was made completely superfluous by
Darwin's theory of natural selection. The adaptations of organisms-Iong
cited as the most conspicuos evidence of intelligent design in the
universe could be explained by purely mechanistic causes. For
evolutionary biologists, the flower of a maglonia has a function not a
purpose. It was not signed in order to propagate the species, much less
to delight us with its beauty but instead came into existence because
magnolias 'with brightly colored flowers reproduced more prolifically than
magnolias with less brightly colored flowers. The unsettling implication of
22
Unit 1 Introduction to Evolutionary Biology
this purely material explanation is that, except in the case of human
behavior, we need not invoke, nor can we find any evidence for, any
design, goal, or purpose anywhere in the natural world.
It must be emphasized that all of science has come to adopt the way of
thought that Darwin applied to biology. Astronomers do not seek the
purpose of cornets or supernova, nor chemists the purpose of hydrogen
bonds. The concept of purpose plays no part in scientific explanation.

Evolution as Fact and Theory

Is evolution a fact, a theory, or a hypothesis? Biologists often speak of the
"theory of evolution," but they usually mean by that something quite different
from what non-scientists understand by that phrase.

You must be aware of the fact that in science, a hypothesis is an informed

conjecture or statement of what might be true. A hypothesis may be poorly
supported, especially at first, but it can gain support, to the point at which it is
effectively a fact. The facts are hypotheses that have acquired so much
supporting evidence that we act as if they were true. A scientific theory is a
mature, coherent body of interconnected statements, based on reasoning and
evidence, that explain a variety of observations. Given these definitions,
evolution is a fact. But the fact of evolution is explained by evolutionary theory.

In ‘The Origin of Species’, Darwin propounded two major hypotheses: that

organisms have descended, with modification, from common ancestors; and
that the chief cause of modification is natural selection acting on hereditary
variation. Darwin provided abundant evidence for descent with modification,
and hundreds of thousands of observations from paleontology, geographic
distributions of species, comparative anatomy, embryology, genetics,
biochemistry, and molecular biology have confirmed this hypothesis since
Darwin's time. Thus the hypothesis of descent with modification from common
ancestors since long had the status of a scientific fact.

The explanation of how modification occurs and how ancestors give rise to
diverse descendants constitutes the theory of evolution. We now know that
Darwin's hypothesis of natural selection on hereditary variation is correct, but
we also know that there are far more causes of evolution than Darwin realized,
and that natural selection and hereditary variation themselves are more
complex than he imagined. A body of ideas about the causes of evolution,
including mutation, recombination, gene flow, isolation, and random genetic
drift, many forms of natural selection, and other factors, constitute our current
theory of evolution, or "evolutionary theory". Like all theories in science, it is
incomplete, for we do not yet know the causes of all of evolution, and some
details may turn out to be wrong. But the main tenets of the theory are well
supported, and most biologists accept them with confidence.
BOX 1.1: Ethics, Religion and Evolution.
In the 'world of science’, the reality of evolution has not been in doubt for more than a
hundred years, but evolution remains an exceedingly controversial subject. The
creationist movement opposes the teaching of evolution in public schools, or at least
demands "equal time" for creationist beliefs. Such opposition arises from the fear that
evolutionary science denies the existence of God, and consequently it denies any
basis for rules of moral or ethical conduct.
Our knowledge of the history and mechanisms of evolution is certainly incompatible
with a literal reading of the creation stories in the Bible's Book of Genesis, as it is
23
Block 1 Evolutionary Biology-I
incompatible with hundreds of other creation myths that people have devised. A
literal reading of some passages in the Bible is also incompatible with physics,
geology, and other natural sciences. But does evolutionary biology deny the
existence of a supernatural being or a human soul? No, because science, induding
evolutionary biology, is silent on such questions. By its very nature, science can
entertain and investigate only hypotheses about material causes that operate with at
least probabilistic regularity: It cannot test hypotheses about supernatural beings or
their intervention in natural events.
Evolutionary biology has provided natural, material causes for the diversification and
adaptation of species, just as the physical sciences did when they explained
earthquakes and eclipses. The steady expansion of the sciences, to be sure, has left
less and less to be explained by the existence of a supernatural creator, but science
can neither deny nor affirm such a being. Indeed, some evolutionary biologists are
devoutly religious, and many nonscientists, including many priests, ministers, and
rabbis, hold both religious beliefs and belief in evolution.
Wherever ethical and moral principles are to be found, it is probably not science, and
surely not in evolutionary biology. Opponents of evolution have charged that
evolution by natural selection justifies the principle that "might makes right," and
certainly more than one dictator or imperialist have invoked the "law"" of natural
selection to justify atrocities. But evolutionary theory cannot provide any such precept
for behavior. Like any other science, it describes how the world is, not how it should
be. The supposition that what is "natural" is "good" is called by philosophers, the
Naturalistic fallacy.
Various animals have evolved behaviors that we give names such as cooperation,
monogamy, competition, infanticide, and the like. It is not a scientific question to
determine whether or not these behaviours are morally right or appropriate. The
natural world is amoral as it lacks morality altogether. Despite this, the concepts of
natural selection and evolutionary progress were taken as a "law of nature" by which
Marx justified class struggle, by which the Social Darwinists of the late eighteenth
and early nineteenth centuries justified economic competition and imperialism, and
by which the biologist Julian Huxley justified humanitarianism. All these ideas are
philosophically undefensible instances of the naturalistic fallacy. Infanticide by lions
and langur monkeys does not justify it in humans, and evolution provides no basis for
hurnan ethics.

1.8 SUMMARY
• Evolution is the unifying theory of the biological sciences. It aims to
discover the history of life and the causes of the diversity and
characteristics of organisms.

• Danwin's evolutionary theory, published in The Origin of Species in

1859, consisted of two major hypotheses: first, that all organisms have
descended, with modification, from common ancestral forms of life, and
second, that a chief agent of modification is natural selection.

• Darwin's hypothesis that all species have descended with modification

from common ancestors is supported by so much evidence that it has
become a well established fact in biology. His theory of natural selection
as the chief cause of evolution was not broadly supported until the
"evolutionary synthesis" that occurred in the 1930s and 1940s.

• The evolutionary theory developed during and since the evolutionary

synthesis consists of a body of principles that explain evolutionary
24 change.
Unit 1 Introduction to Evolutionary Biology
• The implications of Darwin's theory, which revolutionized western
thought, include the ideas that change rather than stasis is the natural
order; that biological phenomena, including those seemingly designed,
can be explained by purely material causes rather than by divine
creation; and that no evidence for purpose or goals can be found in the
living world, other than in human actions.

• Like other sciences, evolutionary biology cannot be used to justify beliefs

about ethics or morality. Nor can it prove or disprove theological issues
such as the existence of a deity.

1.9 TERMINAL QUESTIONS

1. Describe main features of Darwin’s “Theory of Evolution”.

2. What are the fundamental principles of evolution as stated by Modern

Evolutionary Sysnthesis.

3. Discuss the Philosophical Issues Related to Darwin’s Evolutionary

Theory.

1.10 ANSWERS
Self-Assessment Questions
1. a) i) "to unfold or to reveal or manifest hidden potentialities”.

ii) “the change in the properties of group of organisms over the

course of generations”

iii) population

iv) the genetic material

b) A current example of evolution is the trend of antibiotic resistance

development in microbial population. Before the 1940s, the
bacteria infections like tuberculosis and pneumonia had little hope
of being cured. However, by 1965, with the discovery of antibiotic
drugs and subsequent advances in their synthesis led to the
conquest of most bacterial diseases. But at present most bacteria
are resistant to antibiotics like penicillin, ampicilin, that were
supposed to have vanquished them.

2. a) Philosophy of Essentialism states that there existed ideal forms of

everything, including a human. Variation is attributed to
the imperfections manifested in individuals (defects of the
underlying perfect essence/form).

b) The Theory of Special Creation holds that God created each

species uniquely, exactly as it is today.

c) Scala nature represents a gradation from inanimate objects and

barely animate forms of life, through plants and invertebrates, up
through ever "higher" forms of life. Humankind, which is both
physical and spiritual in nature, fanned the link between animals
and angels.
25
Block 1 Evolutionary Biology-I
d) Carlous Linnaeus

e) Lamarck argued that species differ from one another because they
have different needs, and so use certain of their organs and
appendages more than others. The more strongly exercised
organs attract more of the "nervous fluid," which enlarges them,
just as muscles become strengthened by work. Lamarck, like most
people at the time, believed that such alterations, acquired during
an individual's lifetime, are inherited a principle called inheritance
of acquired characteristics.

3. a) Charles Robert Darwin was a clergy at Cambridge University. He

served as a naturalist on a ship (the H.M.S. Beagle) to map the
waters of South America. During his voyage he has made many
observations related to variation in life forms. He assembled all this
thoughts and observation in a book titled “The Origin of Species”.
In the book he gave the theory of descent with modification and
natural selection being the casual agent for the evolution.

b) Thomas Malthus

c) It means that all species, living and extinct, have descended,

without interruption, from one or a few original forms of life.
Species that diverged from a common ancestor were at first very
similar, but accumulated differences over great spans of time, so
that some are now radically different from one another.

d) According to Charles Darwin “Natural Selection” is the cause of

evolution.

4. a) True; b) True; c) True; e) True

f) True; g) True; h) False

5. a) Different

b) Not

c) Genes

d) Allels

e) Populational

f) Natural Selection

Terminal Questions
1) Refer to section 1.4

2) Refer to section 1.6

3) Refer to section 1.7

26
Unit 2 The RNA World and Origin of Life

UNIT 2
THE RNA WORLD AND
ORIGIN OF LIFE

Structure
2.1 Introduction 2.6 Formation of
Macromolecule
Objectives
Iron-Sulfur World
2.2 The Origin of Life
Hypothesis
2.3. Origin of Life: The Big
Clay World
Bang “Origin of the Earth”
Hypothesis/Gene first
2.4 The Chemical Evolution of Hypothesis
Life
Cell-Like structures:
2.5 Origin of Life on Earth Protobiont and Coacervates

Theory of Spontaneous 2.7 RNA World Hypothesis

Generation
2.8 Evolution by Chemical
Disproving Spontaneous Selection
Generation by Louis Pasteur
2.9 The Origin of Genetic Code:
Urey-Miller Experiment Amino Acids as Cofacors in
a RNA World

2.10 Summary

2.11 Terminal Questions

2.12 Answers

2.1 INTRODUCTION
Origin of life is a long-standing and argumentative topic, with various
mechanisms proposed on it. According to one of the proposed theory,
lightning in the early environment with successive formation of amino acids
provided the basic ingredients for the formation by uniting to form long polymer
chains. According to another theory, during the Archean period which is
estimated to be at 4 - 2.5 BYA (billion years ago), a chemical processes took
place at submarine volcanic vents and life originated at those depths which 27
Block 1 Evolutionary Biology-I

was also protected from the ultraviolet radiation that prevailed at the time due
to the lack of an ozone layer. According to the third hypothesized explanation,
life arose from the burning of the carbon and hydrocarbons in comets in the
atmosphere.

It has also been proposed that life may have evolved in intertidal pools that
were persistently drowned and dried out in the sun, a process supported by
the geological record.

According to RNA World theory, there was a point in primitive Earth's history,
about 4 BYA, when the primary living substance was RNA or something
chemically similar to RNA. From last 50 years, this theory has progressed from
discussion to widespread acceptance. In this chapter, we recap the core logic
behind the RNA World and discuss some of the most significant recent
developments made to support and expand on this logic. This chapter also
explain how RNA-mediated molecular collaboration could aid in the genesis
and early evolution of life (Fig. 2.1).

Fig. 2.1: Schematic diagram showing all the prospects about how the life
originated on earth.

Objectives
After studying this unit, you would be able to:
 explain how life originated on earth;

 describe the formation of marcomolecules on earth; and

 understand the hypothesis of the RNA world.

28
Unit 2 The RNA World and Origin of Life

2.2 THE ORIGIN OF LIFE

• The majority of study on the beginning of life does not focus on fossils,
but rather on laboratory studies of chemical events that might have
occurred on Earth 4 BYA.

• If an electric discharge or ultraviolet light is passed through a solution

containing simpler molecules (such as amino acids, sugars, and
nucleotides), many of the molecular building blocks of life (such as
amino acids, sugars, and nucleotides) can be created. Once the basic
building pieces are in place, the creation of a simple replicating molecule
is critical.

• Although we don't know what the first ancestral replicating molecule

was, multiple lines of evidence indicate that RNA came before DNA.
Single-stranded RNA, for example, is simpler than DNA, which is
invariably double-stranded.

• To read or copy nucleotide information, DNA requires enzymes to

"unzip" the two strands. DNA always has the shape of a double helix.
RNA, on the other hand, can interact with its surroundings directly. It can
be immediately read or reproduced.

• RNA can also take on a variety of shapes based on its nucleotide

sequence. RNA will serve as an enzyme (or "ribozyme") in some of
those structural configurations, catalyzing biological reactions. There are
RNA molecules that act as RNA polymerases, catalyzing RNA
replication.

• A self-replicating molecule would be one of the most basic biological

systems imaginable. Other modest lines of evidence show that RNA
preceded DNA, such as "prebiotic soup" studies that obtained the
nucleotide U rather than T (Fig. 2.2).

• The "RNA world" refers to the early stage of life when RNA was the
hereditary molecule. Later in history, life began to use DNA. One reason
for the switch from RNA to DNA could be that RNA-based life was
limited by its relatively high mutation rate.

• Asexual living forms cannot exist if the total harmful mutation rate
exceeds one. HIV and other modern RNA viruses have a mutation rate
of roughly 104 per nucleotide. This reduces their coding ability to
approximately 104 nucleotides, or approximately 10 genes.

• New refined life forms would not evolve unless the mutation rate was
decreased. Because those events occurred on a molecular scale, the
fossil record teaches us little about the genesis of life.

29
Block 1 Evolutionary Biology-I

Fig. 2.2: The picture depicts hydrothermal systems near komatiite shields,
chemotrophs on mid-ocean ridges, and diverse life forms in coastal,
lacustrine, and oceanic waters.

• During initial first few MYA a huge asteroids hit and vaporized the
oceans. The temperatures were too hot to support life. Life could not
have evolved before around 4 billion years ago. The oldest known rocks
are 3.8 billion years old and found at Isua, Greenland.

• Chemical residues in these rocks might have become the chemical

remains of life forms. This type of chemical evidence is inherently
ambiguous because it could have been created by a non-biological
process. Some scientists and geologists regard it as evidence of life, but
few believe it completely.

• The rocks might have undergone too much metamorphism to retain any
fossil cells, assuming that cells existed at that time. Cell fossils were
discovered in a variety of locations between 3 - 3.5 BYA. Until recently,
the earliest fossil cells were assumed to come from 3.5 billion-year-old
rocks from Western Australia's Apex Chert. Other evidence for fossil
cells from the 3 - 3.5 BYA is available. Cells had most likely evolved by
3.5 BYA, or shortly thereafter.

30
Unit 2 The RNA World and Origin of Life

2.3 ORIGIN OF LIFE: THE BIG BANG “ORIGIN OF

THE EARTH”
• The genesis of life is inextricably tied to the formation of the universe
and the birth of our planet. The "Big Bang" idea sheds light on the origins
of the universe.

• The cosmos began as a hot, dense point some 13.8 billion years ago,
quickly expanding and cooling over time. Conditions led to the synthesis
of hydrogen, helium, and a trace of lithium in the first 20 minutes
following the Big Bang.

• This cosmic story includes the origin of Earth. The Earth formed within
the solar system approximately 4.6 billion years ago.

• The development of Earth involved accretion from the solar nebula,

which resulted in the construction of a stony entity that eventually
became our globe.

• Life, on the other hand, appeared considerably later, most likely between
3.5 and 4 billion years ago, in the form of primitive, single-celled
organisms.

• While the "Big Bang" and the beginning of Earth set the stage for life's
potential, the precise methods of life's emergence are still a subject of
scientific investigation.

• Many scientists believe that the Earth, along with other planets, meteors,
asteroids, and comets, was formed at least 4.6 billion years ago.

• According to the protoplanet nebular scenario, the solar system evolved

from a vast cloud of gases and elements created by previously existing
stars.

• Hydrogen and helium were the simplest and most abundant gases, but
other, heavier elements had been produced by nuclear fusion and were
present as well

• The accumulation of particles within this cloud formed a gravitational

attraction, causing other particles to be pushed from the outer borders to
the center.

2.4 THE CHEMICAL EVOLUTION OF LIFE

• The process wherein life originated from non-living or inorganic matter is
termed as chemical evolution through chemical reaction processes.

• This theory posits that the earliest organic substances necessary for life
were borne out of their inorganic counterparts. This was a vital step that
set the platform for life as the cells that are used by living things evolved
at this stage.

• Currently, the theory of chemical evolution suggests that life began a

step-by-step advancement of nonelemental substances. It is speculated 31
Block 1 Evolutionary Biology-I

that this slow and meticulous procedure might have resulted in the
biochemical systems seen in organisms today.

• This idea was first initiated by Aleksandr Pavlovich Oparin from Russia
which was stated in his book “Origin of life on earth” in 1936. The
knowledge of chemical evolution can help in understanding the
phenomenon of abiogenesis.

• The realms of chemistry, biology, and earth’s history form the basis for
this scientific exploration.

2.5 ORIGIN OF LIFE ON EARTH

There are several theories explaining how life originated on earth. Several
projecting theories provide different perspectives on this complex question:

Modern Theory: Suggests that life evolved gradually through chemical stages
in which primitive earth created complex organic substances.

Electric Spark Theory: Implying a random electric spark in the early Earth’s
atmosphere could also be one of the possible and responsible reason for life’s
precursors.

Molecules Meeting on Clay: Postulated that life’s predecessor molecules

developed on clay surfaces, enabling them to combine and synthesize larger
structures.

Deep-Sea Vents Theory: To hydrothermal sites in volcano active areas on

the earth or at sea zones as possible home for genesis of life.

Panspermia Theory: Shows how complex molecules and even primitive life
may have been brought to earth by interplanetary dust, meteoroids, or comets.

2.5.1 Theory of Spontaneous Generation

• The ancient Greek philosopher Aristotle, who lived from 384 to 322 BC
was one of the scholars to propose the concept of generation. According
to this theory life could arise from nonliving matter especially if it
contained a substance called pneuma or "vital heat."

• To support his idea Aristotle pointed out instances where previously

absent animals appeared in environments. For example he mentioned
how fish would suddenly appear in formed puddles of water.

• This belief, in generation continued for centuries and found support in

various observations. For instance proponents of the theory noted that
frogs would appear on the banks of the Nile River during its flooding.
Similarly mice seemed to emerge in barns filled with grain and topped
with roofs.

• Jan Baptista van Helmont even suggested that mice could arise from
rags and wheat kernels left exposed in a container for three weeks under
conditions for mouse populations to flourish.

32
Unit 2 The RNA World and Origin of Life

• However an Italian physician named Francesco Redi challenged this

prevailing notion through an experiment conducted in 1668.

• Redi placed meat into six containers; two left open to the air; two
covered with gauze and two tightly sealed. Only the uncovered jars
developed maggots over time.

• This led Redi to conclude that maggots were formed when flies laid their
eggs on the meat. This finding contradicted the idea of generation (Fig.
2.3)

Fig. 2.3: Francesco Redi's experimental setup included an open container, a

container sealed with a cork top, and a mesh-covered container that
allowed air but not flies in

2.5.2 Disproving Spontaneous Generation by Louis

Pasteur
• The 19th century debate on spontaneous generation engaged scientists
on both sides until the Paris Academy of Sciences intervened, offering a
prize for resolution.

• Louis Pasteur, a renowned French chemist focused on microbial

fermentation, accepted the challenge. In 1858, Pasteur's filtration of air
through a gun-cotton filter revealed microorganisms, challenging the
idea that exposing broth to air introduced a "life force."

• Pasteur designed swan-neck flasks in 1858, boiling broth to sterilize

them (Fig. 2.4). These flasks allowed air exchange but prevented
airborne microorganism entry.

• If a life force other than airborne microorganisms caused microbial

growth, it would access the broth, while microorganisms would not.

• Pasteur correctly predicted that intact swan-neck flasks maintained

sterile broth, but if broken, microorganisms contaminated the flasks,
allowing microbial growth.

33
Block 1 Evolutionary Biology-I

• His experiments definitively disproved spontaneous generation, earning

Pasteur the prestigious Alhumbert Prize in 1862.

• In a subsequent 1864 lecture, Pasteur declared "Omne vivum ex vivo"

("Life only comes from life"), emphasizing the irreparable blow dealt to
the doctrine of spontaneous generation by his swan-neck flask
experiment.

Fig. 2.4: a) Louis Pasteur, a French scientist who finally invalidated the long-
contested idea of spontaneous generation; b) The flasks used in
Pasteur's experiment had a unique swan-neck design that enabled air to
enter but prevented bacterial and fungus spores from entering; c)
Pasteur's experiment was divided into two parts. The soup in the flask
was sterilized by boiling it in the first section. This broth remained
contaminant-free after cooling. The flask was boiled in the second
portion of the experiment, and then the neck was snapped off. This
flask's broth became infected.

2.5.3 Urey-Miller Experiment

• Since the early 19th century, the basic blocks of life have been found and
defined. Two key achievements in the synthesis of molecules of primary
importance to the genesis of life were recognized.

• Adolf Strecker achieved the first laboratory synthesis of an amino acid,

alanine, from acetaldehyde (CH3CHO), ammonia (NH3), and hydrogen
cyanide (HCN) in 1850.
34
Unit 2 The RNA World and Origin of Life

• A few years later, in 1861, Alexandr Butlerov used a strong alkaline

catalyst (NaOH) to achieve the first laboratory synthesis of sugar
combinations from formaldehyde (HCHO). Although these discoveries
were fascinating, they had nothing to do with the origins of life.

• As a result, little progress was made in developing a scientific

description of the genesis of life, which was proposed by Aleksandr
Ivanovich Oparin.

• Oparin pioneered the concept of chemical evolution, which can be

considered as the forerunner of Darwin's theory of evolution. Life,
according to this theory, is the product of a series of spontaneous
chemical reactions that build progressively complex chemical structures.
He proposed that such chemical development would occur within the
primitive Earth's oxidizing atmosphere.

• After a while, he revised his early notions and changed the early
atmosphere to a greatly reducing environment. Similarly, the English
biologist John Haldane, who was the first to introduce the concept of a
"prebiotic soup" where chemical evolution occurred, had similar thoughts
at the same time.

• The concepts of Oparin and Haldane were only presented theoretically.

Stanley Miller and Harold Clayton Urey offered experimental evidence of
the idea of chemical evolution in 1953. They devised and constructed an
experiment to recreate a hypothetical early Earth ecosystem.

• A gaseous combination of hydrogen (H2), methane (CH4), ammonia

(NH3), and water (H2O) was exposed to an electric discharge that
replicated storm lightning in this experiment. The mixture was linked to a
hot bulb filled with liquid water. Within a week, 15% of the carbon
existing as methane had been transformed into other simple carbon
molecules.

• These chemicals included formaldehyde (HCHO) and hydrogen cyanide

(HCN). These compounds were subsequently combined to generate
simple molecules like formic acid (HCOOH) and urea (NH2CONH2), as
well as more complicated molecules with carbon-carbon bonds like
glycine and alanine (Fig. 2.5).

• The resulting creation of a vast number of organic compounds, including

numerous amino acids, was experimental proof of chemical evolution
theory.

• This experiment demonstrated that the chemistry between simple

molecules, which were abundant in the primordial Earth's atmosphere,
resulted in the synthesis of essential chemicals, which may have
contributed to the emergence of life on Earth.

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Fig. 2.5: Urey Miller Experiment.

SAQ 1
a) What Combination of gases was used in Urey miller Experiment?

b) How Life originated according theory of Spontaneous generation?

c) Which scientist disapprove theory of spontaneous generation?

d) How life originated according to protoplanet nebular scenario?

2.6 FORMATION OF MACROMOLECULES

There are several theories for the formation of macromolecules on earth such
as: Iron-Sulfur World Hypothesis; clay world hypothesis or Gene first
hypothesis and RNA world Hypothesis.

2.6.1 Iron-Sulfur World Hypothesis

• Metabolism-first hypothesis suggests that since hydrogen and carbon
dioxide are observed, there is a reversible Krebs cycle occurring within
an iron-nickel-sulfur catalyst.

• A complicated, autocatalytic sequence of metabolic reactions occurs

onto the surface of iron sulfide, generating novel and advanced catalysts
as well as metabolic sequences.

• The metabolic-first scenario or Iron-Sulfur World Hypothesis because of

its dependence on iron sulfide.

• Firstly, it was developed by German chemist Günter Wächtershäuser

who said that the first metabolic reactions started on the surface of
minerals like iron sulfide and nickel sulfide near hydrothermal vents
located at oceans’ bottom.
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Unit 2 The RNA World and Origin of Life

• These minerals are catalysts that turn existing inorganic materials like
carbon monoxide (CO) and/or carbon dioxide (CO2) into carbon.
CO2 is produced by volcanic activities penetrating into oceans’ floor
along ridge systems.

• These compounds undergo the process of releasing carbon with the aid
of hydrogen. Hydrogen become available from the vents and may also
be derived from ocean water containing hydrogen sulphide (H2S).

• The conversion of inorganic to organic molecules was accelerated by the

reducing power (ability to release electrons) of hydrogen sulfide. This
process involves iron sulfide as a simple substrate material on rock
surfaces.

• In addition, the iron sulfide donated electrons to COs, forming carboxyl

acid initiated the reactions that resulted in the formation of long chain of
amino acids that produced proteins.

• As the mixing happened, the energy necessary for these reactions was
supplied by the redox-driven process due to the chemical disruption.

2.6.2 Clay World Hypothesis or Gene First Hypothesis

• There are chances that a kind of mineral, known as clay, might have
contributed to the putting together of the genes and how life started
off. Zircon grains from the oldest known terrestrial material showed that
clays once abounded on earth about 4.4 BYA thus may have been
prevalent when life is thought of to have begun.

• The clay minerals are layered molecular stacks where subsequent

molecules can attach. Therefore, when organic molecules interrelate
with clay, they remain close together in order to undergo reaction and
form long chains of molecule links. Some layers in the clay are made up
of a distinct chemical, elements and compounds.

• Differences in this way could provide the clay with similar properties to
genes.

• Such as charged ions from one layer of clay might act as template for
and catalysed the formation of the next layer of clay. Each new layer is a
replicate of the previous one; any errors occurring while packing alumina
and silica are duplicated in all subsequent copies across all layers.

• These mistakes should serve as effective mutations if they increase in

the efficiency of replication. The generation of such mutations will give
rise to selective advantage by making the replication of new chemicals
faster, thus satisfying our definition about life formation.

• Catalysis by the ions present in the clay layers may also accelerate the
organics in RNA reactions as well as RNA polymerase. Under such
circumstances, it is possible that the clays found in Earth’s seas might
have produced the first auto-replicate organic molecules. This has come
to be known as the clay world hypothesis.
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Block 1 Evolutionary Biology-I

• However, an American chemist Luke Leman, an English chemist Leslie

Orgel and Iranian-American chemist M.Reza Ghadiri noted in 2004 that
it is possible to create polymers within aqueous solution as well. When
present in vascularized tissues or solution, it is possible for carbonyl
sulfide that is a simple gas in volcanic gases and deep-sea-vent
emissions to generate from amino acids. This implies that polymerization
might have arisen in the prebiotic soup.

2.6.3 Cell-Like Structures: Protobiont and Coacervates

• Protobiont refers to an agglomeration of primitive biomolecules and
acromolecules surrounded by an enclosure like lipid bilayer which
separated internal chemical environment from that of the surroundings
around it.

• Abiogenesis involves the study of origins of life arising from non- living
matter and has a crucial role. These are transitional forms leading from
simple organic compounds to primordial life.

• Protocell is defined as being auto-organized, having an internal

organization and involving a cell-bounded envelope or similar boundary.

• Prototype features can grow, reproduce but incorrectly, and can carry
out metabolic functions. Coacervate and microspheres are also included
among the protobionts that have a membrane barrier consisting of
multiple units.

• Liposomes and/or coacervates may have represented an early stage in

the evolution of protobiont.

• Coacervates which means “to assemble together or cluster”, namely

protein, carbohydrate, nucleic acid molecules and other associated
charged polymers are in the state of aggregations surrounded with
water.

• Measuring range of the microscopy is 1 to 100 micrometers under

osmotic effect. Alexander I. Oparin speculated a structure of “organic”
macromolecules with envelope of water molecules. Although it is not a
membrane, this organization of water molecules would have acted as a
physical barrier for these organic molecules against their
surroundings. He called these structures coacervates.

• Evidence of coacervates that exhibit some lifelike traits: They are able to
draw molecules selectively out of surrounding water and build them into
their own structures.

• Chemicals in coacervates also arrange themselves in such a way, they

do not come together as if they were just a random collection of
molecules.

• Some coacervates comprise enzymes, which are used for particular

chemical transformation. It is exemplified by an event where glycogen
phosphorylase in coacervate was offered with glucose-1-phosphate and
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Unit 2 The RNA World and Origin of Life

starch synthesized which fused with coacervate and wall and resulted in
its expansion.

• With addition of enzyme amylase, the starch converted to maltose that

got out of coacervates.

• However, coacervates were not considered as living structures since

they comprise fluid mass lacking a definite membrane. Nevertheless,
under favorable circumstances, coacervates can enlarge their sizes and
even break down into small cells.

• Another possibility is that the early cell-like structure which was termed
as ‘Microspheres’ was made up of a collection of organic
macromolecules with a double-layered outer wall.

• A portion of the protein material forms a double-boundary structure that

encloses the microsphere. Although these barriers lack lipids, they
exhibit membrane-like properties and resemble the shape of a cellular
membrane.

• Depending on the osmotic potential in the surrounding solution,

microspheres swell or shrink. They also exhibit internal movement
(streaming) comparable to that of live cells and contain chemicals that
operate as enzymes. Microspheres can guide the production of
polypeptides and nucleic acids by using ATP as a source of energy.

• They have the ability to absorb material from their surroundings and
produce buds, resulting in a second generation of microspheres.

2.7 RNA WORLD HYPOTHESIS

• According to the RNA World hypothesis, catalytic RNA molecules were a
transitional form between nonliving materials and the first cells. Sidney
Altman and Thomas Cech shared the Nobel Prize in Chemistry in 1989
for discovering ribozyme.

• According to the RNA World hypothesis, the primordial form was an

RNA-based living system that later developed into the life-forms we see
today, in which DNA stores biological information and proteins exhibit
this information. RNA was a living entity distinguished from nonliving
systems by its ability to store and transmit information (genotype) as well
as express that information (phenotype).

• Evolution requires the ability to record and modify heritable information,

as well as a sorting process that distinguishes beneficial modifications
from harmful ones.

• The former is a feature of genotype, whereas the latter occurs as a result

of phenotypic diversity among individuals. These are some of the
qualities of RNA. Most phenotypes involve the creation and dissolution
of phosphodiester links in RNA or DNA (Fig. 2.6).

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Block 1 Evolutionary Biology-I

Fig. 2.6: All the aspects of RNA World Hyothesis (Higgs and Lehman, 2015).

• The chemistry of these events is exactly what is required for nucleic acid
replication. This finding lends credence to the hypothesis of a primordial
RNA World in which RNA is responsible for recreating itself.

• If an RNA molecule could replicate itself while allowing for mistakes or

mutations, it would display many of the features of current life and so be
deemed alive.

• According to the theory, the first living entities on earth used RNA as
both genetic material and for catalytic activity, which includes genetic
material replication. The first living creatures, according to the RNA
World hypothesis, were made up of three components: a ribozyme with
RNA polymerize activity (the synthesis of a complex RNA molecule by
combining smaller and simpler molecules), a template RNA to drive
polymerization, and a physical container (membrane).

• Two RNA molecules are required to initiate this process and for the
ribozymes to catalyze reactions. To act as a template for the synthesis of
a new RNA molecule, an RNA molecule must be unfolded and exposed
to the monomer (simpler molecule) that will polymerize on it (Fig. 2.7).

Fig. 2.7: Diagrammatic illustration of RNA World Hypothesis.

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Unit 2 The RNA World and Origin of Life

• The production of two ribozymes is critical in this scenario. A container is

also required to keep the genetic material and the molecules it encodes
together. Without this, the material would dissipate and interact with
nothing.

• To conclude, according to the RNA World hypothesis, our ancestors

began as two RNA molecules (a self-replicating ribozyme) within a lipid
membrane. Nucleotide monomers "leaked" into the membrane and
polymerized into new ribozyme copies.

• This process proceeded, producing more RNA molecules and increasing

the molecular weight of the lipid membrane. According to the RNA World
hypothesis, catalytic RNA molecules were a transitional form between
nonliving materials and the first cells. The RNA World concept is based
on the knowledge that RNA can have both a genotype and a phenotype.

• RNA is involved in protein synthesis machinery; for example, ribosomal

RNA catalyzes protein synthesis. Ribonucleoside triphosphates, such as
ATP and GTP, are the basic units of cellular energy. These compounds
are involved in nearly every energy-transfer function in all organisms and
are even components of electrontransfer cofactors like NAD
(nicotinamide adenine dinucleotide), FAD (flavin adenine dinucleotide),
and SAM (S-adenosyl methionine).

• The genotype is the fundamental sequence of nucleotides along the

RNA, similar to how a modern organism's genotype is the sequence of
nucleotides along the DNA in the chromosome. Catalytic RNA, for
example, has between 30 to 1,000 ribonucleotides in its main sequence,
which determines its genotype. Cech and colleagues found the
Tetrahymena ribozyme, which spans 400 nucleotides from the head (the
5' end) to the tail (the 3' end). This RNA is an intron (an intervening
segment between two genes) that divides two ribosomal RNA-coding
sections of the Tetrahymena genome.

• The folded state of ribozymes can have an active region that allows the
RNA to catalyze a chemical reaction on a substrate, similar to a protein
enzyme.

• After ten generations, the average RNA in the population had enhanced
by a factor of 30 in cleaving DNA substrates and attaching one of the
resultant fragments to its own 3' end.

• Individual ribozymes with mutations at positions 94, 215, 313, and 314
demonstrated a catalytic efficiency (ability to pick up a new 3' tail utilizing
DNA substrates) 100 times more than the previous ancestral sequence.
This experiment proved that RNA molecules in solution can have live
organism-like characteristics that allow them to evolve. Each RNA has a
survival (substrate catalysis) and reproduction (ability to be reverse- and
forward-transcribed) function.

• The RNA World theory is based on the idea that RNA predates the
usage of proteins for most biological functions. The showing that RNA
can copy itself is thus the missing piece of evidence.

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Block 1 Evolutionary Biology-I

• An RNA with a replicate phenotype would be required whether the RNA

World used only one type of self-replicating RNA or a suite of interacting
RNAs.

• For example, David Bartel and colleagues employed test-tube evolution

to look for ribozymes capable of RNA synthesis by looking at which RNA
(from a pool of long RNA strands) can bind the most numbers of the
same short RNA strand.

2.8 EVOLUTION BY CHEMICAL SELECTION

• Chemical selection happens when a chemical in a mixture has unique
features or benefits that cause it to outnumber other compounds in the
mixture.

• Chemical evolution occurs when a population of molecules evolves over

time to form a new population with a different chemical composition.

• Protobionts containing RNA exhibited life-like qualities due to RNA

genomes that were copied and maintained via the catalytic action of
RNA molecules. Further mutations in these RNA molecules could have
resulted in the addition of other new functionalities that are selected by
nature, a process known as chemical selection.

• Chemical selection would have resulted in an increase in complexity in

these protobionts, with RNA molecules gaining capabilities such as the
capacity to link amino acids together to form proteins and other catalytic
functions.

2.9 THE ORIGIN OF GENETIC CODE: AMINO

ACIDS AS COFACTORS IN A RNA WORLD
• In this hypothesis it was proposed that aminoacyl-RNAs for Glu, Asp,
Lys, Tyr, His, Arg, Cys, and Ser were first chosen as cartridge-type
subunits of three-subunit ribozymes. The proto P-site of the large subunit
of ribozymes accommodated aminoacyl-RNA subunits acting as
cofactors.

• It is also expected that there will be no direct connotation between amino

acids and codon and anticodon pairs. Duplication of the proto P-site
could have resulted in the proto A-site, allowing multi-subunit ribozymes
to interact with two-cartridge-type aminoacyl-RNA subunits at the same
time.

• The insertion of two cartridges at random would have immediately

stopped the enzymatic activity of multi-subunit ribozymes.

• If two tandemly aligned pairs of codons and anticodons specified two

cartridges, dozens of different active pockets in multi-subunit ribozymes
would have rapidly emerged, resulting in the development of extant
species' metabolic pathways.

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Unit 2 The RNA World and Origin of Life

• The powerful driving force of Darwinian selection outlined here could

have resulted in the creation of the fundamental genetic code for
catalytic amino acids (Fig. 2.8).

Fig. 2.8: Origin of Genetic Code.

SAQ 2
a) What are protobionts?

b) What are coservates?

c) What is the Basic Unit of energy in RNA Synthesis?

2.10 SUMMARY
The RNA World paradigm is being explored, and new discoveries in synthetic
organic chemistry and biology are routinely providing fresh insights. Prebiotic
chemistry is increasingly identifying phenomena that provide solutions to
several (rather than single) problems at the same time. The question of how
RNAs generated copies of themselves (that is, how they replicated) is central
to RNA world study. There are presently multiple putative mechanisms for this
process, with a growing emphasis on those that exhibit autocatalytic feedback.
Cooperation among different molecules was most likely an important
component of the RNA world, and at least three types of molecular
cooperation could have been at work during the genesis of life. Chemical
alternatives to RNA may have existed at some point in Earth's history, and
several efforts are being made to locate and assess such structures. Another
process that had a significant impact on the organization of the living state,
from small molecules to giant molecules and cell-like structures, was network
formation. This unit explains that how life originated on earth with focus on
various theories like Big bang theory, theory of spontaneous generation; Urey
miller Experiment and the origin of the genetic code in RNA world. The
production of two ribozymes is critical in this scenario. A container is also
required to keep the genetic material and the molecules it encodes together.
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Block 1 Evolutionary Biology-I

Without this, the material would dissipate and interact with nothing. To
conclude the RNA world hypothesis, our ancestors began as two RNA
molecules within a lipid membrane. Nucleotide monomers "leaked" into the
membrane and polymerized into new ribozyme copies.

2.11 TERMINAL QUESTIONS

1. Write a brief description about origin of life on earth.

2. Write a brief note about Gene First Hypothesis.

3. Write a brief note RNA world hypotheis.

2.12 ANSWERS
Self-Assessment Questions
1. a) A gaseous combination of hydrogen (H2), methane (CH4),
ammonia (NH3), and water (H2O) was exposed to an electric
discharge that replicated storm lightning in Urey Miller experiment.

b) According to theory of spontaneous generation life could arise

from nonliving matter especially if it contained a substance called
pneuma or "vital heat."

c) Louis Pasteur

d) According to the protoplanet nebular scenario, the solar system

evolved from a vast cloud of gases and elements created by
previously existing stars.

2. a) Protobiont refers to an agglomeration of primitive biomolecules

and acromolecules surrounded by an enclosure like lipid bilayer
which separated internal chemical environment from that of its
surroundings.

b) Coacervates, namely protein, carbohydrate, nucleic acid

molecules and other associated charged polymers are in the state
of aggregations surrounded with water.

c) ATP and GTP

Terminal Questions
1. Refer to Section 2.5.

2. Refer to Subsection 2.6.2.

3. Refer to Section 2.7.

Acknowledgment
Fig. 2.4: a, b) : https://bio.libretexts.org/Bookshelves/Microbiology/Microbiology_
(OpenStax)/03%3A_The_Cell/3.01%3A_Spontaneous_Generation

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