Darwin’s Theory of

Evolution by Natural Selection

A new theory of biological species

Waseda University, SILS,

Introduction to History and Philosophy of Science
The Facts about Evolution

In the early modern period, due to colonialism and empire

building, European naturalists, working in centralized
botanical gardens and national zoos, investigated an
unprecedented variety of animal and plant specimens. Starting
in the 18th century, naturalists began to systematically
investigate the fossil remains of various organisms and
compare these with living organisms.
In the early half of the 19th century, it became clear that there
had once existed entire families of flora and fauna (plants and
animals) that had passed out of existence, and that moreover, in
the periods – that is, geological strata – in which these creatures
existed, much of the flora and fauna that are alive today did not
exist.
The evidence for large-scale biological change was gathered
slowly and was still ongoing when Darwin was working.

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Various Theories of Evolution

Although there was a lot of disagreement about how these

changes had taken place, and what they implied, by Darwin’s
time, most naturalists accepted that there had been some
changes in biological species.
However, even if we accept that there has been change in
species throughout the history of the earth, we might have
several different theories about how this change occurred.
All of the theories advanced before Darwin argued for some
kind of directed change – in some sense responding to, and
hence directly influenced by, the environment and the actions of
organisms.
Darwin tried to distinguish his theories from these by arguing
that evolutionary changes were based only on naturally
occurring processes – processes that are still occurring around us
now.
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The Theory of Evolution by Natural Selection

The theory of evolution by natural selection is a theory about the

mechanism by which evolution occurred in the past, and is still
occurring now. The basic theory was developed by both Darwin
and Wallace, however, Darwin gave a much fuller argument.
The theory states that biological change takes place with two
basic characteristics:
1) Variation: Random variations occur in the traits of individual
organisms and are passed on to their offspring.
2) Struggle for existence: There is an existential competition
that insures advantageous traits are preserved and
disadvantageous traits are eliminated.

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Local Circumstances, Local Advantages
A key result of evolution by natural selection is that it forces us
to look at various binaries such as advantage and disadvantage,
superior and inferior from a local perspective.
This means that there cannot be traits that are better in any
absolute sense – there can only be traits that are better suited
for a particular set of circumstances.
§ In an anoxic environment, being able to consume sulfates
is more useful than, say, general intelligence.
Within a species, individuals specialize at exploiting the
environment in various ways, and over long periods this
specialization creates divergence.
§ A group of birds with longer beaks might prefer worms,
while a group with stronger beaks might prefer nuts.
§ Being attractive to, and interacting with, mates of the
opposite sex is an exploitation of local circumstances.
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The Place of Homo sapiens
One of the most controversial aspects of the theory of evolution
is the implications that it has for human beings.
According to current thinking, there were a number of other
species (or varieties) of the homo genus that were closely related
to humans, but which are now all extinct (Homo habilis, H.
erectus, H. ergaster, H. heidelbergensis, H. neanderthalensis, H.
floresiensis, etc.). H. habilis lived some 2.3 million years ago,
whereas the Neanderthals died out about 24,000 years ago. The
whole genus is thought to be about 2.5 million years old and to
have diverged from the pan genus of great apes about 5 million
years ago.

Key Point
Humans did not evolve from apes of the pan genus. Both pan
apes and humans evolved from now extinct common ancestors.
Moreover, we are still part of the ape family (Hominidae).
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The Ape Superfamily

The term homo sapiens was coined by Carl Linnaeus in 1758.

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Evolutionary Theories in the 19th Century
In the 19th century, there were many conflicting ideas about
evolution and the meaning of the existence of variations among
organisms. Many natural philosophers, believed that God had
created a fixed number of species and that since that time there
had been hybridization and mixing (variation), but no
fundamental change.
In the 1790s, Darwin’s own grandfather, Erasmus, had
published poems putting forward the idea of evolution.
In the 1810s, Jean-Baptiste Lamarck outlined a comprehensive
theory of evolution in which animals, individually, become more
complex and better adapted and then pass these changes onto
their offspring. This kind of theory is known as Lamarkism.
In 1844, Vestiges of the Natural History of Creation (Robert
Chambers) presented a natural history in which all forms were
in transformation and everything develops from previous
forms.
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19th Century Geology: Neptunism
In the 18th and 19th centuries the empirical study of rocks and
minerals was of increasing economic importance. The earliest
theories of rock formations were developed in the mining
schools in the German states.
Abraham Werner (1749–1817), a professor at the mining school
in Freiburg, set out a theory of stratification, in which he
claimed that different types of rocks were laid down in different
periods of the earth’s history by the gradual drying of a
primordial ocean.
In the early 19th century, this was combined with a hypothesis
of cataclysmic changes and developed as a theory that helped
geologists prospect for different types of rocks and minerals.
There were also conservative thinkers, who linked the
primordial ocean of neptunism with Noah’s flood mentioned in
the Bible, but this was not the majority.
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19th Century Geology: Plutonism
In the 19th century, it became clear that the same kinds of rocks
had been formed in different periods of the earth’s history, so
geologists began to study the fossils embedded in the strata. It
became clear that there were the same kinds of fossils at
comparable levels all over the earth.
Geologist like James Hutton (1726–1797) and Charles Lyell
(1797–1875) began to focus on the role of volcanoes and argued
that the earth had a molten core – which was called plutonism.
They argued that geological formations are not the result of
cataclysmic changes in the past, but the long-term gradual
result of processes we see around us now. This became known
as uniformitarianism.
Hutton:
“We find no vestige of a beginning,—no prospect of an end.”

Naturalists began to contemplate vast stretches of time.

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W. Buckland, Geology and Mineralogy …

…Considered with Reference to Natural Theology, 1837

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Malthus’ Social Ideas

§ Beginning in the end of the 18th century, the rising

industrialism in Britain created a massive growth of urban
poor. This lead to the rise of poverty as a social problem
and to various attempts to address it.
§ In 1798, Thomas Malthus (1766–1834) published his Essay
on the Principle of Population.
§ Malthus argued that human population increases
geometrically, while food supplies increase arithmetically.
§ This means that at some point, population growth will
always outstrip food supplies and there will be a struggle
for existence, especially among the poor.
§ Thus, Mathus saw the conditions of poverty, which had been
created by the Industrial Revolution, as a necessary fact of the
human condition.
§ This was a thoroughly pessimistic doctrine.
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Charles Darwin (1809–1882)

§ 5th child of a wealthy, agnostic doctor.

§ Started in medicine, then graduated
from Cambridge to be a pastor, but felt
no sense of vocation.
§ Took the opportunity to pay his own
way as a naturalist on a surveying
expedition: 5 years on HMS Beagle
§ Became a respected member of the
British scientific elite.
§ After 20 years, published his ideas
about evolution.
§ Was very often sick with various
psychosomatic disorders.
§ Worked endlessly on scientific projects
until the day he died.
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The Voyage of the Beagle
§ While on the voyage of the Beagle, Darwin read Lyell’s
Principles of Geology.
§ He collected thousands of specimens of plants and animals
and carried out extensive studies of the geography of
South America and various Pacific islands.
§ He came to believe that the Andes were formed over long
periods of time, by the landmass slowly rising and the seas
receding.
§ He became convinced that the atolls of Pacific islands were
produced by the coral growing as the mountains sank.
§ In Tierra Del Fuego, he observed native people living in
the wild and became convinced that humans are not so far
removed from animals and show great diversity amongst
themselves.
§ In the Galapagos Islands, he observed considerable
diversity but he collected erratically and did not even tag
by island.
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Secret Notebooks

When he returned to England, he sent

out all of his specimens to be cataloged
by experts.
§ John Gould realized that all of the
different types of birds that Darwin
had collected on the Galapagos were
various species of finches.
Darwin began to keep a series of secret
notebooks in which he wrote his ideas
about the “transmutation” of species.
From the beginning, he was interested in
the origin of life and the human species.
During this time, he read Malthus’ Essay.
Darwin’s first sketch of his This gave Darwin the idea for a
concept of speciation mechanism for evolution.

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Becoming Respectable
After returning from the voyage of the Beagle, Darwin began to
publish his ideas about the geography of South America,
handbooks of the flora and fauna of the places he visited and
an account of his travels.
He made a name for himself as a natualist and became a friend
and colleague of many of the most important biologists in
England.
Occasionally, he would try to interest someone in his ideas
about transmutation, but no one was convinced.
Because Darwin was himself a member of the social elite, he
tried to distance himself from other evolutionists, such as
Lamarck and Chambers. Also, his wife was very religious.
§ At this time, in Britain, evolution was regarded as a
dangerous idea, associated with France, the revolution,
and social and political unrest.
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The Origin of Species, 1859–1872
When Darwin realized that the established scientists were not
interested in his ideas, he began to cultivate the friendship of
younger men who were on the rise such as Thomas Huxley
(1825–1895) and Joseph Hooker (1817–1911). Over a period of
many years, through correspondence, he tried to convince them
of his ideas.
In the 1850s, he also started a correspondence with Alfred
Wallace (1823–1913). This lead to Wallace sending Darwin a
short paper in which he independently described a theory of
evolution by natural selection.
It was decided by Lyell and Hooker, who both knew of
Darwin’s work, that the two men should make a joint
presentation at the Linnean Society acknowledging their
independent discovery. No one took any notice of this.
Darwin then spent over a year developing the argument which
was published as On the Origin of Species, 1859.
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The Origin of Species: Variation (Chaps. 1 & 2)

Chap. 1: Darwin pointed out that animal breeders artificially

select preferred characteristics from domestic populations
(pigeons, cows, dogs, etc.) in which there is a lot of random
variation.
Over many generation these small differences produce very
different types of animals (varieties of the same species).

Chap. 2: Darwin argued that a similar selection occurs in nature.

He then began to discuss the nature of species. He pointed out
that it is often hard to demarcate between species; it is hard to
decide what are different species as opposed to different
varieties.
He argued that we could think of well-marked varieties as
“incipient species” – that is, newly forming, or emerging.

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The Origin of Species: Struggle for Existence (Chap. 3)

Chap. 3: Darwin argued that individual organisms and species

are all in a struggle for existence.
§ This was contrary to the prevailing view – that nature was
harmonious and showed evidence of a beneficent creator –
a doctrine known as natural theology.
Instead, Darwin claimed that life is a complex struggle – not
always dramatically violent, as in predators and prey, but
usually more subtle, as a plant struggling against drought, or
the introduction of new species into a habitat, etc.
He argued that this struggle then resulted in what he called
“natural selection.” In the 6th edition of the book, he adopted
the terminology of Herbert Spencer and referred to the
“survival of the fittest.”

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The Struggle for Existence

The Origin of Species, 1872 (6th edition)

“A struggle for existence inevitably follows from the high rate
at which all organic beings tend to increase. Every being…must
suffer destruction during some period of its life…otherwise, on
the principle of geometrical increase, its numbers would
quickly become so inordinately great that no country could
support the product. Hence, as more individuals are produced
than can possibly survive, there must in every case be a
struggle for existence, either one individual with another of the
same species, or with the individuals of distinct species, or with
the physical conditions of life. It is the doctrine of Malthus
applied with manifold force to the whole animal and vegetable
kingdoms; for in this case there can be no artificial increase of
food…”

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The Origin of Species: Natural Selection (Chap. 4)

Chap. 4: In this chapter, Darwin explains the core of his theory:

§ Nature varies, randomly.
§ Some of variations are more advantageous than others.
§ Because there is a struggle for existence, any advantage
will be important, however slight.
§ In the long run, those organisms that have a slight innate
advantage will survive more often than those that do not,
passing on their advantages to their offspring.
Through this mechanism, Darwin argued that a population
will change by the accumulation of small, but favorable
advantages, over vast periods of time.
Again, he referred to artificial selection, calling natural selection
“unconscious artificial selection.” But notice that nature does
not chose preferred traits in the way that a breeder does.

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Natural Selection

The Origin of Species, 1868 (5th edition), Introduction

“Owing to this struggle [for existence], variations, however
slight and from whatever cause proceeding, if they be in any
degree profitable to the individuals of a species, in their
infinitely complex relations to other organic beings and to their
physical conditions of life, will tend to the preservation of such
individuals, and will generally be inherited by the offspring.
The offspring, also, will thus have a better chance of surviving,
for, of the many individuals of any species which are
periodically born, but a small number can survive. I have called
this principle, by which each slight variation, if useful, is
preserved, by the term Natural Selection, in order to mark its
relation to man’s power of selection. But the expression often
used by Mr. Herbert Spencer of the Survival of the Fittest is
more accurate, and is sometimes equally convenient.”

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Darwin’s concept of divergence

The diagram represents divergence against time, as earth strata

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The Initial Reaction

The Origin was a very popular book, and went through six
editions from 1859 to 1872. Darwin tapped into the thinking of
the time and pitched evolution as a form of progress similar to
the progress of the English nation that his contemporaries
perceived. England was no longer as conservative as it had
been when he was young and a belief in social progress using
science and industry fit well with the evolutionary idea of
biological progress.
Literal interpretations of the Bible were coming under
increasing criticism and many were prepared to be convinced
of the fact of biological evolution.
Darwin’s close supporters, such as Huxley, Hooker and Lyell,
were influential and helped Darwin carry out the social aspect
of the argument for evolution.

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The Reaction to Natural Selection
Although the Origin convinced many naturalists of the fact of
evolution, many people remained skeptical about the theory of
natural selection. People raised the following objections:
§ It is random and open-ended.
§ Is not a goal directed process.
§ Where the goal was implicitly understood as white,
Victorian, upper-class males.
§ It does not imply directed progress.
§ This was a challenge to the establishment and to the
mercantile values of the rising middle class.
§ Is not an open system, as in Lamarck’s theory. That is,
individuals cannot improve themselves throughout the
course of their lives and pass on these improvements.
Most naturalists of the 1870s and 80s became evolutionists, and
even called themselves “Darwinists,” but did not accept natural
selection, which was the core of Darwin’s theory.
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Darwinists without Natural Selection

Even many people who

supported Darwin, did not
accept, or understand, natural
selection.
For example, Herbert Spencer,
used Darwin’s ideas to explain
human progress towards fitter
individuals – a very
progressivist view of evolution.
Ernst Haeckel, an avid
Darwinist, thought that
evolution unfolded according
to a plan, like embryonic
development (“ontogeny
recapitulates phylogeny”).
Haeckel’s “Tree of Life”
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The Religious Reaction
Darwin’s ideas gave a profound shock to the religious
establishment, and to the naturalists who supported it with the
theory of natural theology. Darwin’s theory was perceived as
being much more difficult to reconcile with a creative, all
powerful god, than those of Copernicus, Galileo or Newton.
If we take natural selection seriously it is difficult to see how a
god could have created us, and hence it is difficult to see how
we could have any special place in a god’s creation.
There were a number of public disputes between religious
authorities, older naturalists and the Darwinists.
§ Bishop Wilberforce and Huxley at the British Association
for the Advancement of Science meeting.
§ There was an ongoing debate in newspapers and journals.
A number of naturalists argued for “theistic evolution,” and
even many of Darwin’s colleagues, such as Lyell and Wallace,
could not fully accept the natural selection of human beings.
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The Modern Synthesis
In the 1930s–40s, there was a revival of the theory of evolution
through the mechanism of natural selection. It was argued that
the new science of genetics made it clear that mutations in
genes could be completely random and that these could then be
adaptively selected for by purely environmental pressures.
The modern synthesis brought together all the different
branches of life sciences and viewed their findings through the
light of natural selection.

Dobzhanzky, 1973:
“Nothing in biology makes sense except in the light of
evolution.”
There was a renewed interest in the evolution of the human
race and in thinking about the implications of evolution
through natural selection as a basis for understanding our
social and moral place in the world.
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Natural Selection and Human Nature
Some of the most serious implications of the theory of natural
selection are on our understanding of human nature.
If human beings evolved from a primate ancestor, it might mean
that even our most “noble” traits – such as our ability to love,
our feeling of the divine, our creativity, our love of knowledge,
our concern for others, etc. – are not the gifts of a beneficent
god, but are rather the chance results of natural processes.
While it is easy to see that our base traits – such as greed, lust,
selfishness, etc. – would have helped our ancestors to survive, it
has been more difficult to see this with our noble traits.
We are still in the process of developing models of behavior
that show how these traits would have been advantageous to
our ancestors. That is, scientists are currently still articulating
the paradigm of natural selection to cover more and more
phenomena.
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Overview
Over the course of his life, Darwin went from being a devout
Anglican, who had intended to become a minister and believed
that the natural world was evidence of God’s providence, to
being an agnostic who believed that human beings were the
chance result of natural processes.
The revolution in thought and worldview brought about by the
theory of natural selection is one of the most profound to have
effected us.
Although almost all biologists now accept natural selection, it is
still difficult for many lay people to accept. It is still disputed
from the perspectives of both religion and “common sense.”
From the perspective of worldviews, we should remember that
it often takes a long time for a new worldview to take hold. A
change in worldview involves many cultural and social factors
that do not always follow a clear logic.
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