• enumerate the scientist and cite their respective contributions in the

development of evolutionary thought

• describe Jean Baptiste Lamarck’s hypothesis on evolutionary change

• discuss Charles Darwin’s theory of evolution by natural selection

• explain the Modern Synthesis as the unified theory of evolution

17th Century: John Ray: the "species" concept

The first scientist (in the modern sense of the word) to carry out a thorough
study of the natural world was the Englishman John Ray (1627 - 1705). Ray
was a brilliant student who, unusually for his time, did not take holy orders
on completing his degree at Cambridge (largely due to the social and
religious upheavals associated with the Civil War, but also because of his
own personal beliefs). Forced to resign his Fellowship at the university, he
was sponsored by his friend Francis Willughby (1635 - 1672), who shared Ray's
scientific interests, to develop his catalogues of the living world. Ray's
particular interests lay with plants, for which he developed an early
classification system based on physiology and anatomy. During this work Ray
established the modern concept of a species, noting that organisms of one
species do not interbreed with members of another, and used it as the basic
unit of taxonomy.

Ray also studied fossils, recognising them as having formed from once-living
organisms, and grappled with the contradictions between the Biblical account
of creation and the evidence of change and extinction that he saw in his
fossils. Deeply religious, he rejected any possibility of an old and changing
Earth, as did all scholars of his time.

18th Century: Carl Linnaeus & the modern taxonomic system

Ray's ideas on taxonomy were picked up and extended by the better-known Carl
Linnaeus (1707 - 1778). He was fascinated by plants, paying botany much more
attention than was required for his medical studies at university, and took
up the new idea that plants reproduced sexually, using differences in
reproductive structures to develop a system for classifying plants. He moved
on to study animals, and to help make sense of the huge volume of data
accumulated during his teaching and research gave all his specimens a
descriptive Latin binomial, or two-word, name. Linnaeus used these names
systematically in his classification system, which he published as "Systema
naturae". The "Systema" built on Ray's earlier work and catalogued the
diversity of living things in a cohesive and logical manner - the now-
familiar hierarchical way of arranging organisms, from the all-inclusive
Kingdoms through Classes and Orders to the Genus and Species of each living
thing. Linnaeus went so far as to include humans in his system, and believed
that humans and the great apes were so closely related that they should be
placed in the same genus. However, he didn't actually do so, to avoid
contradicting church teachings.
Although a religious man who believed that all species were created together,
Linnaeus still gave some thought to the apparent age of the Earth. Fossils
were now well-accepted as the remains of past creatures, sometimes found far
from the sea, and he was uncertain that this distribution could have been
achieved in the time provided by the Biblical flood. He was not alone in
this, as contemporary scholars of both science and history were beginning to
question the calculations of Archbishop Ussher that gave the age of the Earth
as 6000 years.

Buffon on evolution and the age of the Earth

The Frenchman Georges Louis Leclerc, Comte de Buffon (1707 - 1788), was both
intellectually brilliant and extremely hard working (though he viewed himself
as naturally lazy) and produced an enormous body of work over his lifetime.
He set out the current knowledge of the whole of natural history in the 44-
volume "Natural History" ("Histoire Naturelle"), a series that greatly
increased popular interest in science. He also contributed to the debate
over the age of the Earth (begun by Isaac Newton), suggesting that our planet
had formed in a molten state and that its gradual cooling must have taken
far longer than the 6000 years Ussher and other theologians allowed (or the
50,000 permitted by Newton). In fact, unlike previous scientists, Buffon
attempted to answer this question by experimental means, something that has
been described as "a landmark event in science" (John Gribbin, 2002).

Buffon also gave consideration to the concept of evolution - a concept that

was in circulation long before Charles Darwin provided a mechanism by which
evolution could occur - wondering about the role of vestigial organs, which
appeared to give the lie to the idea that creation achieved perfection of
form, and about the possibility of species descending from earlier ancestors.

Erasmus Darwin's thoughts on evolution

Charles was not the first in his family to consider the concept of evolution.
His grandfather Erasmus Darwin (1731 - 1802) was a successful country doctor
who published widely in many scientific fields. He was also a poet, and his
book "The Loves of the Plants" introduced the public to the intricacies of
plant taxonomy and reproduction. Another book, "Zoonomia", set out Erasmus'
ideas on evolution. He was aware that modern species were different to fossil
types, and also saw how plant and animal breeders used artificial selection
to enhance their products. He knew that offspring inherited features from
their parents, and went so far as to say that life on Earth could be descended
from a common ancestor. While Erasmus believed in the original creation of
life, his God was "hands-off" from that point on. His ideas were not widely
accepted in England, but in France Lamarck was developing similar views of
evolutionary change.
Cuvier's contribution to palaeontology

Georges Cuvier (1769 - 1832) was interested in biology from childhood, an

interest that he developed further while living in the French countryside
during the Revolution. He read both Linnaeus and Buffon and worked on his
own ideas on classification and taxonomy, before joining the Museum of
Natural History in Paris, studying and writing on comparative anatomy. His
work was extremely useful in interpreting the remains of fossil animals and
relating them to living species. Cuvier also classified animals according to
their body plan (as vertebrates, molluscs, those with jointed exoskeletons
and those with radial symmetry), a major advance in thinking about
relationships. His extensive studies of fossils gave rise to the science of
palaeontology, and he recognised that particular groups of fossil organisms
were associated with certain rock strata. This last finding meant that it
became possible to place the strata into order by relative age of the fossils

The catastrophism model of Earth's history

Cuvier's palaeontological studies told him that large numbers had become
extinct. To explain this, he used the concept of catastrophism: a series of
catastrophes, one of which was recorded in the Biblical story of the flood,
had caused repeated waves of extinction. Areas were then repopulated by
migration from unaffected areas: there was no room in this model for the
evolution of new species. In his view, life had existed unchanged on Earth
for hundreds of thousands of years, ever since the Creation. Cuvier's
adherence to the concept that species were "fixed" and unchanging meant that
he rejected the model of evolution developed by his fellow Frenchman,
Lamarck.

Lamarck's concepts of evolution and inheritance

Jean-Baptiste Lamarck (1744 - 1829) also worked at the Natural History Museum
in Paris, but his views on species were the opposite of Cuvier's. His model
of evolution proposed that individuals were able to pass to their offspring
characteristics acquired during their own lifetimes. (At the time this was
a perfectly acceptable model of change, given that nothing at all was known
about the processes of inheritance.) But what annoyed Cuvier was Lamarck's
proposal that species did not go extinct, but instead evolved into another
form. In fact, Lamarck went further, stating that evolution produced more
complex organisms from simple ancestors, and that this process of change
took time.

Etienne Geoffroy Saint-Hillaire (1772 - 1844) elaborated on Lamarck's views.

Like Lamarck, he felt that the environment could produce changes in living
things, but went on to suggest that if these changes were harmful, then the
organism would die; only those well-adapted to the environment would survive.
This is a foretaste of Darwin's theory of natural selection, but Geoffroy
never went on to develop his idea further. This is because both his
suggestions, and Lamarck's ideas about inheritance of acquired
characteristics, were thoroughly ridiculed by Cuvier. And since Cuvier was
such a prominent scientist, his attacks carried a lot of weight. Most
scientists accepted the principle of catastrophism that he championed so
strongly, until the work of Englishmen James Hutton and Charles Lyell.

James Hutton and the principle of uniformitarianism

James Hutton (1726 - 1797) made a significant contribution to the

understanding of the geological processes that shaped the Earth. He was a
keen chemist but also developed a strong interest in geology. Hutton
recognised that the Earth was extremely old. He saw that there was no need
for global catastrophes to shape the surface of the Earth. Instead, given
sufficient time, the gradual ongoing processes of erosion, sedimentation,
and uplift could produce the geological features he saw. This concept became
known as the principle of uniformitarianism.

Without the concept of an extremely old, and slowly changing, Earth, Darwin
would not have had the time available for his model of evolution to work. In
fact, Darwin specifically applied Hutton's concept of gradual change, or
gradualism, to his model of how species evolved.

18th century: Charles Lyell

Charles Lyell (1797 - 1875) went to Oxford to study mathematics and law but
turned to geology after being introduced to Hutton's work. He met Gideon
Mantell, who had discovered several different dinosaurs in English rocks,
and this led him to the serious study of geological history. Lyell travelled
widely in Europe, where he observed ancient raised seabeds separated by lava
flows, and became convinced that Hutton's model of gradual geological change
was correct. He collected a large amount of supporting evidence for
uniformitarianism and set this out in the "Principles of Geology", a book
that had a tremendous influence on Darwin. As well as building on the idea
of gradual long-term natural changes as the shaping force of the Earth's
surface, Lyell considered the origins of plants and animals. While he
believed in the special creation of all species now in existence, he also
recognised that many species had become extinct and been replaced by others.

Charles Darwin and the theory of evolution by natural selection

Charles Robert Darwin (1809 - 1882) was one of six children born to Robert
& Susannah Darwin. Robert was a well-respected local doctor and also
something of a private investment banker; the family was always very well
off. Charles was fascinated by science, particularly natural history, from
a young age. His father wished him to become a doctor, but the traumatic
experience of observing an operation on a non-anaesthetised child caused
Charles to reject that career and he became took classes in geology and
natural history, particularly marine biology.

Robert Darwin still wished his son to have a career and so arranged for him
to study for the clergy at Cambridge. Many country clergymen managed to
combine their priestly duties with an interest in natural history, so this
seemed the obvious thing for Charles to do.

However, Charles once more ignored his official studies and took classes
reflecting his interest in the natural world, including botany and geology.
His teachers in these courses viewed Darwin as an outstanding and hardworking
pupil, but his father was still set on him becoming a country parson.

The summer after he graduated from Cambridge, Charles received a letter that
was to change his life. His botany professor, John Henslow, had put his name
forward to join the crew of HMS Beagle, on a surveying expedition to South
America. Contrary to popular myth, this was not the naturalist's position
(which was filled by the ship's doctor). Instead, the Beagle's captain,
Robert FitzRoy, required a "gentleman companion" to provide company and
conversation on the voyage. FitzRoy found Darwin acceptable and the Beagle
left on what was to be a five-year voyage on December 27, 1831. Darwin was
yet to turn 23.

Robert Darwin (who was paying Charles' expenses) expected that his son would
settle down on the voyage and come home ready to take up a country parish.
His university tutors and scientific friends had different expectations:
Charles was to collect scientific specimens and send them back to England.
This sort of amateur collecting was a significant hobby for the upper classes
in Darwin's time - and provided a livelihood for less well-off men such as
Alfred Russel Wallace, who was later to provide the impetus for the
publication of "On the Origin of Species".

Darwin's thinking was enormously influenced by the work done by previous

scientists. Not least of these was Charles Lyell. Darwin took Lyell's
"Principles of Geology" for reading matter on the voyage. What he read, and
later confirmed at first hand in South America, led him to accept the
uniformitarian approach to Earth's history. This is significant because it
allowed for the vast age of the Earth, necessary for his model of gradual
evolutionary change.

Darwin travelled extensively in South America while the Beagle continued its
surveying duties. He made extensive fossil collections and noticed that these
fossils were found in regions now occupied by their slightly different
descendants. This led him to think about factors affecting a species'
distribution. He also found evidence supporting Lyell's theory of gradual
geological change, such as fossils and ancient sea beds now far from the
sea, and witnessed first-hand how a large earthquake could raise the land.

He also visited the Galapagos Islands and noted how the finches and iguanas
there resembled those of the South American mainland. Contrary to the usual
story, "Darwin's finches" did not provide him with a "eureka!" moment. He
noticed how the islands' giant tortoises varied from island to island, but
the significance of the varied finch species did not strike him until after
his return to England.

Darwin was greeted with considerable scientific acclaim on returning home.

This was due to the quality and quantity of the scientific specimens he
brought with him, and in fact he first made his name as a geologist, not a
biologist. Robert Darwin became resigned to the fact that his son was going
to follow his own path, and Charles settled down to writing about his travels,
and studying his specimens. And he began to develop his theory of evolution
by natural selection as a coherent explanation for his observations on the
form and distribution of species, tying it into the concepts developed by
other thinkers such as Lyell, Lamarck and Malthus. While he quickly produced
an outline of this theory, Darwin was to spend the next 25 years refining it
and amassing still more supporting evidence. It took a letter from Alfred
Russel Wallace to push him into publication.

Alfred Russel Wallace arrives independently at a theory of evolution

Alfred Russel Wallace (1823 - 1913) did not have the same advantages in life
as Charles Darwin. Largely self-taught, he had always had an interest in
natural history but not the funds to indulge it. Unhappy with what was
essentially a dead-end job he managed to save enough money to fund a trip to
South America, with the intention of collecting specimens and selling them
to wealthy private collectors. This was a difficult life but Wallace was
moderately successful. More importantly, he also attended various scientific
meetings, published papers, and began corresponding with Darwin.

While on a major expedition to South-East Asia Wallace began to give serious

consideration to how the species he was observing might have evolved. Like
Darwin, he was influenced by the ideas on limits to population size developed
by Malthus. Quite independently of Darwin, Wallace came up with the idea
that the best-adapted organisms in a population would survive to breed,
passing on their adaptations to their offspring. He worked this insight up
into a paper that he sent to Darwin, asking for his comments and assistance
in getting it published.

The theory of evolution by natural selection

Darwin was shocked to receive Wallace's paper. He had been sitting on his
theory for 25 years and here was another naturalist coming up with the same
concept. Darwin felt that Wallace should have priority in publication, but
was persuaded by his friends that he should produce a pr�cis of his own
work. The two documents were read together at a meeting of the Royal Society,
but it is Darwin's contribution that we remember today. "On the Origin of
Species by Means of Natural Selection, or the preservation of favoured races
in the struggle for life" was published in 1859.

The following statements represent the heart of the theory of how species
evolve by the process of natural selection, developed by Darwin and Wallace:

 Observation 1: organisms of all species can produce so many offspring

that their population size would increase exponentially if all
individuals that are born reproduce successfully.
 Observation 2: populations tend to remain stable in size, except for
seasonal fluctuations.
 Observation 3: environmental resources are limited.
 Inference 1:production of more individuals than the environment can
support leads to a struggle for existence among individuals of a
population, with only a fraction of offspring surviving each generation.
 Observation 4: individuals of a population vary extensively in their
characteristics; no two individuals are exactly alike.
 Observation 5: much of this variation is heritable.
 Inference 2: survival in the struggle for existence is not random, but
depends in part on the hereditary make-up of the surviving individuals.
Those individuals whose inherited characteristics best fit them to their
environment are likely to leave more offspring than less-fit
individuals.
 Inference 3: this unequal ability of individuals to survive and
reproduce will lead to a gradual change in a population, with favourable
characteristics accumulating over the generations.

Or, to put it another way:

1. Natural selection = differential reproductive success.

2. Natural selection occurs as a result of interaction between the
environment and genetic variability in the population.
3. The outcome of natural selection is the adaptation of populations to
their environment.

The modern synthesis

A major problem for Darwin lay in the apparent lack of a mechanism by which
features could be inherited. Gregor Mendel had developed his theory of
heredity at much the same time as Darwin was grappling with the theory of
evolution by natural selection. However, although Mendel published his
findings, it seems that Darwin never read the paper. In fact, the
significance of Mendel's work wasn't properly grasped until early in the
20th century.

The "rediscovery" of Mendel's research led to the understanding of how

various characteristics, coded for in an individual's DNA, could be passed
on to successive generations. However, both Mendel and the early geneticists
recognised only the classic "either/or" states of discrete characteristics.
It seemed there was no way to explain how natural selection could operate on
more subtle variations within a population.

It took the development of population genetics to reconcile Mendelism and

Darwinism. As the name suggests, population genetics examines genetic
variation within populations, not individuals, and allows an understanding
of the relationship between this variation and natural selection.

The modern synthesis, a "comprehensive theory of evolution" (Campbell &

Reece, 2002), was developed in the 1940s. It brings together information
from the fields of population genetics, palaeontology, taxonomy,
biogeography, and molecular biology. It "emphasises the importance of
populations as the units of evolution, the central role of natural selection
as the most important mechanism of evolution, and the idea of gradualism to
explain how large changes can evolve as an accumulation of small changes
occurring over long periods of time" (Campbell & Reece, 2002: 446). While
modern evolutionary biology may see healthy debate over some of the
assumptions made by the modern synthesis, the underlying tenets of evolution
set out by Charles Darwin remain secure.