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Natural Selection

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N
Natural Selection
Elena Racevska
Department of Social Sciences, Faculty of
Humanities and Social Sciences, Oxford Brookes
University, Oxford, UK
Natural selection is a process by which organisms
that are better adapted to specific pressures of their
environment tend to survive longer and produce
more offspring, thus ensuring the preservation and
multiplication of those favorable traits through
generations, at the expense of the less advanta-
geous ones. It is one of the key mechanisms of
evolution, which affects organisms in every
aspect, from their physiology and morphology,
to behavior and ecology. The term was popular-
ized by Charles Darwin (Darwin 1859).
Natural selection has been incorrectly called
“survival of the fittest,” but rather than to the
survival, it refers to a differential reproductive
success of organisms who do and do not carry
beneficial traits that affect how many descendants
they will leave.
The Maturation of the Idea of Natural
Selection
The idea of species changing over time long pre-
dates Darwin’s theory of evolution by natural
selection, and it is possible to follow its
# Springer International Publishing AG, part of Springer Nature 2018
J. Vonk, T. K. Shackelford (eds.), Encyclopedia of Animal Cognition and Behavior,
https://doi.org/10.1007/978-3-319-47829-6_542-1
development back to the ancient Greece. Anaxi-
mander suggested that all life forms, including
human ancestors, came from water. Empedocles
believed that both animals and plants were formed
from disconnected parts that would come together
in different combinations. Those combinations
that were the most adapted survived, while the
strange ones became extinct. Plato believed in
essentialism and proposed a theory of Forms, or
theory of Ideas, which argued that non-physical
forms (ideas) represent the most accurate reality
(Ross 1951). Aristotle created scala naturae (also
known as the Ladder of Life or the great chain of
being), which he used to explain the relationships
between all living organisms, placing them on this
scale hierarchically, with regard to their structural
and functional complexity. He believed that all
organisms were designed for a purpose, thus
rejecting Empedocles’ view of all organisms orig-
inating by chance (Charlton 1983).
While Middle Ages philosophy emphasized
the role of divine guidance in the creation and
development of life, renaissance naturalists
maintained that this development was mechanical
(Bowler 1989). In the Age of Enlightenment,
these ideas were further developed. Carl Lin-
naeus, a Swedish botanist and taxonomist who
came up with the taxonomic system of binomial
nomenclature, believed that God created a small
number of species, which then interbred and pro-
duced hybrids that became new species (Bowler
1989). Comte de Buffon, a French naturalist,
rejected Linnaeus’ ideas, proposing that species
2 Natural Selection
and even individuals within them differed. He
defined a species as a group of animals that were
able to produce fertile offspring. He was also the
first to suggest a possibility of all animals
descending from a single breeding pair. James
Hutton, Scottish naturalist and the father of mod-
ern geology, advocated uniformitarianism – the
assumption that the laws of the universe that oper-
ate at present are stable across time and space – for
all living organisms, suggesting natural selection
as a mechanism that may be affecting them. He
believed that species formed varieties, some of
which made them better adapted to particular
environments, distinguishing between heritable
and nonheritable variations. Traits that are passed
down from parents to offspring are referred to as
heritable traits. On the other hand, nonheritable
or acquired traits are obtained through individ-
ual’s actions (e.g., developing large muscles or
higher flexibility through exercise) or are a result
of learning (e.g., certain food type preferences or
seeking a particular hiding place to avoid a
predator).
Erasmus Darwin, an English physician and
Charles Darwin’s grandfather, suggested that all
living organisms, from plants to animals, have a
common origin. He believed that a “living fila-
ment” is the cause of all organic life. This “great
first cause,” as he called it, granted the organisms
the power to acquire new parts and improve their
inherent activity over generations (Darwin 1794).
His most important publication is Zoonomia (Part
I published in 1794, Part II published in 1796),
wherein he explains how species should be spread
through the proliferation of the strongest and most
active animals in order to improve. This view is
today regarded as the survival of the fittest.
In 1809, a French biologist Jean-Baptiste
Lamarck proposed a theory of the transmutation
of species. Lamarck believed that living organ-
isms were continuously created by spontaneous
generation, and that their increasing complexity
was prompted by an innate life force. He did not
believe that all organisms shared a common
ancestor, but he recognized the adaptedness of
species to their environment. However, he pre-
sumed that the organisms changed depending on
the organs they used or disused, and these changes
would be passed down to the next generation,
producing adaptations to the environment.
This idea that organisms can pass on the acquired
characteristics to their offspring is known as
Lamarckism or Lamarckian inheritance, or soft
inheritance.
In 1813, William Charles Wells read a paper
before the Royal Society in which he explained
the principle of natural selection in humans.
Studying how different breeds of production ani-
mals are bred through an artificial, human-
orchestrated selection of the best individuals, he
realized that an equally efficient, albeit a slower,
process is carried out by nature on humans. Wells
was interested in the emergence of human races.
He proposed that they arose from accidental vari-
eties and those that were best suited to a particular
country’s climate and diseases became the most
prevalent (Wells 1818). Charles Darwin and
Alfred Russel Wallace were not aware of Wells’
work at the time of publishing On the Origin of
Species, but Darwin later acknowledged Wells for
being the first to recognize the principle of natural
selection. He, however, also recognized the limits
of Wells’ conclusions, as he only noted the natural
selection in humans, and only in respect to race.
Edward Blyth, an English zoologist, also
discussed artificial selection of production ani-
mals, as well as recognized the within-species
variation in nature, but he did not see it as a
potential for formation of new species, but rather
as a way to preserve the one that already exists
(Blyth 1835). He believed that natural selection
operates on within-species variations by remov-
ing those that were less suited for the species’
natural habitat than the “original” form, which
was without modifications. He also did not
believe in a shared ancestry of species.
In 1844, Robert Chambers, a Scottish pub-
lisher and author, anonymously published Ves-
tiges of the Natural History of Creation. Herein
he argued a similar view of evolution to that
proposed by Lamarck. Chambers proposed a cos-
mic theory of transmutation, suggesting that all
currently existing forms, from the solar system to
all living organisms, including humans, have
developed from earlier forms (Chambers 1844).
He argued that life originated by spontaneous
Natural Selection 3

generation and that the reason behind extinction On the Tendency of Species to Form Varieties;
lies in flawed designs – however, a supernatural And on the Perpetuation of Varieties and
force is, according to Chambers, unnecessary. The Species by Natural Means of Selection
book was received with praise, and even Charles
Darwin believed that its publication may have “Extract from an Unpublished Work on Species”
facilitated the acceptance of his “Origin of spe- (by Charles Darwin)
cies,” due to bringing attention to the subject of This essay contains a portion of a few chapters
evolution and removing prejudice. Alfred Russel (“On the Variation of Organic Beings in a State of
Wallace found the book ingenious, but in need of Nature,” “On the natural means of selection,” and
more evidence. “On the comparison of domestic races and true
species”) later published in his On the Origin of
Species. Darwin began his essay with a reference
to De Candolle’s argument of all nature being at
The Theory of Evolution Through
war, “one organism with another, or with external
Natural Selection
nature” (p.46). Darwin agreed, stating, however,
that this war is not a constantly ongoing one, but
The theory of evolution through natural selection
instead recurs at short periods (to a lesser degree)
was independently developed by two British
and occasionally (to a more severe degree). He
naturalists – Charles Darwin and Alfred Russel
further explained that it is also a result of seasonal
Wallace. Wallace believed that every species
differences in climate and food abundance. Spe-
emerged from a closely related one that had
cies differ in their favoring of particular seasons,
already existed. Unaware of Darwin’s at the time
which Darwin saw as an application of Malthus’s
still unpublished work, he autonomously reached
doctrine “with a tenfold force”. He believed that
similar conclusions. He wrote them in an essay,
mankind could double in size more quickly with
which he mailed to Darwin, asking his opinion.
more available resources; however, since animals
His effort resulted in the jointly written publica-
do not have these “artificial means,” the average
tion entitled On the tendency of species to form
amount of their food is constant, but their increase
varieties; and on the perpetuation of varieties and
will in most cases be enormous. If unexpected
species by natural means of selection (Darwin and
factors – for instance, drought – are at play, it is
Wallace 1858). The two scientists presented their
possible that some species will perish, while
work to the Linnean Society of London on 1 July
others will flourish. Darwin explained that this
1858, which was organized by Joseph Dalton
higher than normal increase is likely a result of
Hooker and Charles Lyell. At the time, Darwin
more organisms surviving to the age of reproduc-
had not yet written his ideas as a paper, so he read
tion, and reproducing. Once the environment of
parts of his drafts of the On the Origin of Species,
the flourishing species returned to normal, the
which he had already began writing, as well as
numbers would decrease. However, Darwin pro-
parts of his correspondence with Professor Gray.
posed that this decrease would still occur regard-
Asa Gray, who Darwin met in 1839, was one of
less once the environment was completely full and
the most notable botanists of the nineteenth cen-
could not support more individuals.
tury. Darwin considered him a kindred spirit.
Noting that most environmental changes tend
Wallace’s and Darwin’s presentation was
to be to a fairly small degree, and therefore caus-
published as a science paper on 20 August 1858.
ing only a small change in the inhabitants, Darwin
This publication was the first revelation of their
drew the attention to an example in which the
theory of evolution by natural selection. The fol-
population is small and isolated from others on
lowing year, Darwin published his world-
an island, with their secluded environment under-
renowned book On the Origin of Species.
going a progressing change in a way that its
4 Natural Selection
original inhabitants would soon no longer be per-
fectly adapted to it. Since all individuals have to
struggle for resources, and against their conspe-
cifics, even the smallest variation of a trait that
puts them at advantage could ensure success. If
this variation is heritable, the offspring of the
individuals carrying the variation would also
have better chances. If the species continued to
breed, and if the more suitable characteristics were
selected for over a number of generations, it will
result in a similar change observed previously in
domestic animals after the identical principle of
selection.
In addition to the “natural means of selection,”
Darwin recognized another similar effect – the
battle of the males for the females. He believed,
however, that this selection was less rigorous, as it
does not require the death of the less successful
individual: it just gives them fewer offspring.
Darwin noted that this struggle usually occurs at
times of food abundance, and it does not have an
impact on traits related to food acquisition or
predator avoidance. Instead, it affects secondary
sexual characteristics, therefore carrying implica-
tions for intrasexual competition. These mecha-
nisms comprise sexual selection.
“Abstract of a Letter from C. Darwin to Prof. Asa
Gray” (by Charles Darwin)
Darwin’s essay is followed by the abstract of the
letter he wrote to Professor Gray. Darwin wrote
about a human-orchestrated selection of useful
animals. He believed that, regardless of the degree
of intention with which such selection was carried
out, it was the main force behind the production of
domestic animals. He explained to Gray how
selection only works through the accumulation
of variations and argues that humans have adapted
particular races within a species for different pur-
poses. Darwin also wrote about how the main
reasons for children not completely resembling
their parents are “the changed conditions of exis-
tence” (p. 51). He explained that natural selection
exclusively selected for the good traits. He backed
his claim by saying that any species could, given
enough time, cover the entire earth, if it was not
for the “check” that occurs during some part of
organism’s life or during a shortly recurrent
generation. He explained that this is why only a
small number of organisms born each year survive
long enough to propagate their species.
If a change in the environment occurs, it will
likely cause a variation of the inhabitants. While
some will die out, others will be exposed to the
“mutual action” of a different set of inhabitants
(those remaining in the population). Darwin
explained to Gray his belief that the latter is of a
much higher importance to each individual’s life
than an environmental change. He also stated that
variation can be accumulated in any part of organ-
ism’s structure, as well as useful during any part of
its life. This abstract of Darwin’s letter also
includes an explanation of how the occurrence of
new varieties, new subspecies, or even new spe-
cies (i.e., speciation) will typically exterminate the
previous, less well-fitted parent species. Darwin
believed his letter to be very imperfect, inviting
Professor Gray to use his imagination to fill up the
“very wide” blanks.
Darwin and Gray continued an extensive cor-
respondence. Gray originally did not believe in
the concept of simpler species becoming more
complex over time and even tried to convince
Darwin that all life forms had an inherent, unmis-
takable design. Gray later wrote a collection of
essays entitled “Darwiniana,” which were
published in 1888. Herein, he defended the theory
of evolution, reconciling it with theology by stat-
ing that the two are not mutually exclusive. His
other work focused on the morphological similar-
ities between eastern Asian and eastern North
American plants, a phenomenon that is now
known as the “Asa Gray disjunction.”
“On the Tendency of Varieties to Depart
Indefinitely from the Original Type” (by Alfred
Russel Wallace)
Wallace began his essay by stating the, at the time,
generally accepted belief that the varieties, both in
domesticated species and in the wildlife, are typ-
ically unstable and over time show a tendency to
return to the “normal form” of the species. In his
part of the paper, Wallace aimed to show that the
opposite was true: His belief was that in the nature
many varieties will survive, causing successive
variations progressively departing from the
Natural Selection
original type, while in domesticated animals the
tendency will be to return to the parent form.
Similarly to Darwin, Wallace believed that
wildlife struggle to survive and reproduce, and
those processes require them to fully utilize all
their capacity. The main challenges the wild ani-
mals face come from environmental conditions,
which affect not only the survival of individuals
within a population but of entire species. Wallace
further explained that large animals will not be as
abundant as small ones, and that carnivores will
be less numerous than herbivores. He proposed
that the prevalent view of his time that species’
abundance is directly caused by their fecundity is
not true. Wallace believed that entire world’s pop-
ulation of animals is, despite the fluctuations,
stationary or possibly decreasing by the influence
of humans. He also suggested that every species
reaches its maximum population size within just a
few years of its emergence. For a population to
stay stable in size, equal numbers of organisms
that are born also die. If every parent pair breeds
two offspring, twice the number of the new born
organisms must die. In his opinion, large broods
were, for this very reason, superfluous, as the
majority of the offspring are likely to either
become prey, or die due to environmental condi-
tions (i.e., hunger or harsh climate). Wallace
suggested that a species can increase its popula-
tion size rapidly if there is a constant supply of
wholesome food. Some species can tackle the
problem of this condition not being met by migrat-
ing. Those that are unable to do so will, in
Wallace’s view, remain scarce. The organisms
that die are the weakest and the least perfect.
Wallace thought that all variations have a “def-
inite effect, however slight, on the habits or capac-
ities” of an individual (p.57). Furthermore, if a
variation offered advantages, it will over time
become superior, occurring in the majority of the
population. In addition, he recognized that some
variations would never reach that level of preva-
lence. The advantageous variation will over time
completely replace the original form, and the new
species will be better adapted to the environment.
The new form would not be able to return to its
original, as the organisms carrying the original
5
form would never be able to compete for exis-
tence. The new form would instead continue to
emerge new varieties, the superior of which would
again follow the same principle of replacing the
currently prevalent form. Wallace referred to this
as a progression and continued divergence. He
also noted that variations that arise in aspects of
no perceivable effect on the life preservation do
not follow the same patterns. They can instead
coexist, progressively diverge, or return to the
previous form.
Unlike wild animals, domesticated animals do
not have to struggle for survival and have every-
thing provided for them. Potentially advantageous
variation that occurs in these species is therefore
useless and could exist without the animal’s
awareness. All variations emerging in domesti-
cated animals have an equal chance of being
passed on to the offspring, until humans artifi-
cially select for the preferred traits. Many of the
artificially selected traits would very likely never
be selected for in nature due to them being disad-
vantageous and thus a step towards “inferior
forms” (p.60). Wallace provided the examples of
these, in his opinion abnormal and irregular ani-
mals, in dog breeds and quickly-fattening pigs. If
domesticated animals turn wild, they will there-
fore either return to the original form of the spe-
cies or die out. Because the selection mechanisms
worked so very differently among wildlife and
domesticated animals, Wallace believed that very
little can be inferred about variation in the nature
from simply observing domestic animals.
In the last parts of his essay, Wallace
commented on the Lamarck’s theory of evolution.
He believed that his hypothesis was unnecessary,
as similar results are achieved by the constantly
occurring variation, and those variations that
ensure the most success will survive. He saw this
as proof of the nature’s tendency for continued
progress of the varieties that successively distin-
guish the species from its origins, while the oppo-
site mechanism is at play in domestic animals,
which show a tendency to revert to the original
form. Although these changes occur in small
steps, they are continually “checked” and
balanced out.
6 Natural Selection

The Mechanism of Natural Selection, as Inheritance

Proposed in On the Origin of Species Inheritance is the process of genetic information
According to Charles Darwin’s conclusions being passed on from parent to offspring.
published in the On the Origin of Species, natural Darwin’s hypothetical mechanism of heredity
selection is the inheritance of genetic traits that are was pangenesis. He presented the idea of pangen-
more suitable to the environment. The process is, esis in his 1868 book The Variation of Animals
however, not driven by superiority of one organ- and Plants under Domestication (Darwin 1868).
ism over another in an absolute sense but by a This hypothesis was originally conceived by two
comparative advantage that certain individuals ancient Greek philosophers, Hippocrates and
have over their conspecifics. Natural selection is Democritus. They postulated that an entire organ-
based on four principles: variation, inheritance, ism of a parent participates in heredity. Darwin’s
population growth, and differential survival and theory was almost identical to theirs. He believed
reproduction. that tiny heredity particles, which he called gem-
mules, were transmitted from parent to offspring
(Holterhoff 2014). These gemmules were tiny
Variation
contributions from every tissue and organ of the
Variation in traits is the precondition of natural
parents, collected in their gametes, and transmit-
selection. According to Darwin, organisms do not
ted to the offspring. They were responsible for the
adapt to the environment – the variation in their
configuration of offspring’s every tissue and
traits is a preexisting state, and the environment
organ. Since Darwin regarded gemmules as sim-
simply favors organisms with certain variants of
ilar to plant spores and seeds, he believed it was
these traits. The favored traits enable the individ-
possible for some variations to be expressed by
uals who possess them to survive and reproduce in
the majority of offspring, while others could
greater numbers, thus passing them on to the next
remain dormant for generations before being
generation. Over time, this process produces new
expressed. Darwin offered an explanation of one
species. While some characteristics can be highly
parent’s characteristics dominating over those of
variable (e.g., body size or hair color), others
the other’s, considering this to be the result of that
show very little to no variation – for example,
parent’s gemmules being more numerous or vig-
the number of eyes or limbs among animals of
orous. However, the theory failed to account for
the same species.
the preservation of variations through genera-
Darwin was an externalist and believed that
tions, as this type of inheritance would rapidly
variation was generated by environmental
reduce the original variation to the average of the
changes, which were its necessary prerequisite
original (parents’) traits (Liu 2008).
(Winther 2000). He also proposed natural selec-
Not unlike the majority of the nineteenth-
tion as an external mechanism for adaptation.
century scientists, Darwin was convinced in the
Darwin suggested two different ways in which
idea of blending inheritance. This theory posited
the environment can cause variation: indirectly –
that traits from each parent were averaged and the
through an altered embryonic development, or a
genetic composition of offspring was a uniform,
malfunction of parents’ reproductive systems,
intermediate blend of their parents’. One of the
causing variation in the subsequent generation,
problems with this theory was the lack of expla-
or directly – through an action on the organism’s
nation for the propagation of beneficial mutations:
body through the use and disuse of its organs, with
according to the blending inheritance theory,
the variation displayed in the same generation
favorable, new traits would be unable to spread
(Darwin 1859). In addition, Darwin offered two
through a population since they would be blended
internal mechanisms for variation: crossing
out in the subsequent generation.
between organisms of the same species and cross-
After Darwin’s 1868 publication, Francis Gal-
ing between organisms of different species, which
ton, an English statistician, conducted a series of
he referred to as hybridism.
experiments involving blood transfusion between
Natural Selection
a common lop-eared rabbit and a silver-grey rab-
bit. His experiments negated the pangenesis the-
ory (Galton 1871). Darwin reacted by explaining
that his theory does not state that gemmules occur
are conveyed by blood. Despite this, Galton’s
results were the main reason why the pangenesis
theory was long disregarded (Liu 2008). How-
ever, the experiments of blood transfusion carried
out in the 1950s and 1970 on poultry by Sopikov,
Gromov, and many other Soviet researchers
yielded positive results. A possible explanation
is that heritable changes are easier to detect in
poultry than in rabbits, as it can take many gener-
ations for a trait to be expressed in rabbits. Exper-
iments using parabiosis – surgical joining of
anatomical parts of two animals involving a
blood transfusion – that Boriachok-Nizhnik
(1951) conducted on Angora rabbits and Flanders
rabbits resulted in the so-called vegetative hybrids
of the two species, confirming that blood transfu-
sion alters hereditary traits. Later studies
suggested that another important role may be
that of DNA. Experiments on ducks revealed
that vegetative hybridization was possible using
erythrocytic DNA (Benoit et al. 1960), and the
treated parent was affected as well. It has since
been shown that DNA integration is a mechanism
of horizontal gene transfer through blood transfu-
sion. A proposition that gemmules could include
RNAs, circulating DNAs, and other mobile ele-
ments has been broadly accepted since the discov-
ery of circulating DNA and RNA molecules, as
well as prions in plant sap and in the blood plasma
and serum of both living and deceased animals.
One of the biggest critiques of the pangenesis
theory is its Lamarckian hypothesis that allowed
for the heredity of acquired traits. Darwin
discussed the effects of use and disuse, drawing
a comparison between production animals and
wildlife, suggesting that some changes in traits
that were commonly attributed to these processes
could be results of natural selection. He related the
processes that occur in the nature to artificial
selection or selective breeding of domestic ani-
mals and plants, aimed towards a development of
particular (desired) traits.
7
Population Growth
Darwin and Wallace believed that natural selec-
tion is dependent on a high-population growth.
Darwin originally thought that a population will
increase until its size is in accordance with the
available resources, at which point it will become
stable. However, he refined his ideas after reading
an essay published in 1798 by an economist
Thomas Robert Malthus entitled the “Essay on
the Principle of Population.” Malthus’s central
premise was that a growth of population will
always overpower the growth of food supplies,
thus creating perpetual states of struggle, hunger,
and disease. Human population, is left unre-
strained, would increase beyond their means and
struggle to survive. Darwin and Wallace extended
this incurable struggle for survival to the evolu-
tionary scheme, as they realized that animals and
plants are likely to experience the same pressures.
They proposed that the organisms that were better
equipped to survive will thrive and pass those
more desirable traits on to the following genera-
tion, while the organisms that were less able to
cope with the inherent inequality of the environ-
ment would die out. As offspring numbers are
sometimes higher than what could be supported
by the local environment, they must compete.
This often unconscious competition for resources
will cause the mortality of less-fit offspring. Traits
that ensure more successful coping with environ-
mental challenges (e.g., harsh winters or
droughts) will become more common in the next
generation, which will result in an ongoing pro-
gression of the species. Darwin believed that this
competition for resources will be the harshest
between closely related forms sharing environ-
mental niches, a concept he anticipated.
Differential Survival and Reproduction
Variation occurs among any population causing
individual differences in traits. Some of the vari-
ants can enhance individual’s chances of survival
and reproduction. Individuals who possess traits
that enable them to prevail in the struggle for
resources will contribute more offspring to the
next generation. If these traits are heritable, these
8 Natural Selection
beneficial effects will be reflected in a differential
reproduction – there will be a higher proportion of
organisms carrying those traits in the following
generation. In some cases, the survival or repro-
ductive advantages will be very slight. However,
over many generations, they will nevertheless
become predominant in the population. Even
though natural selection is without a purpose,
traits that carry reproductive advantages will be
selected for.
Types of Selection
There are different types of selection in regard to
the effect they have on traits: directional selection
favors a single extreme phenotype, stabilizing
selection favors an intermediate phenotype over
the extremes, and a disruptive selection favors
extreme phenotypes over an intermediate and
can be a precursor of speciation. If selection acts
to remove the genetic variation from the popula-
tion, it is considered a negative selection (also
known as a purifying selection). If, however,
selection acts to maintain the genetic variation, it
is referred to as balancing selection. Selection that
acts to increase chances of survival is known as
viability selection (or survival selection), while
that which increases organisms reproduction
rates is called fecundity selection (or fecundity or
reproductive selection).
Types of selection that act on an individual
organism are jointly referred to as individual
selection, while gene selection acts on the genes.
Examples of gene selection include kin selection
(an evolutionary strategy that favors the reproduc-
tive success of organism’s relatives, even at a cost
to the organism’s own survival and reproduction)
and the intragenomic conflict (a situation when
genes inside a genome not being transmitted by
the same rules, due to a single gene causing its
own transmission to the detriment of the rest of the
genome). Selection that acts on groups of organ-
isms is called group selection, but its mechanisms
are not yet fully understood. For a group selection
to take place, the trait or behavior that is being
selected for needs to spread out through the entire
group. Group selection has mostly been proposed
in humans and eusocial species (mainly Hyme-
noptera). Finally, with regard to the resource that
the organisms are competing for, selection be
classified as sexual selection (resulting from
mate competition) or ecological selection (also
known as environmental or survival selection, of
asexual selection), which refers to strictly ecolog-
ical processes operating on species’ inherited
traits.
Factors that reduce survival or reproductive
success within a population are called selection
pressures or evolutionary pressures. If they
decrease the occurrence of a trait, they are called
negative selection pressures, while those that
increase the proportion of a trait are called positive
selection pressures. Some selection pressures are
dependent on the density of a population (for
examples, predators, resource availability, supply
of nutrients, or spread of pathogens and diseases),
while others are not (e.g., weather conditions,
changes in temperature or carbon dioxide levels,
or natural disasters).
Alternatives of the Theory of Evolution
by Natural Selection
While the majority of Wallace’s and Darwin’s
contemporaries were relatively promptly per-
suaded by their theory of evolution, few of them
accepted natural selection as the prime driving
force of evolution. The late nineteenth century
brought forward a couple of alternatives to natural
selection.
Theistic evolution, also known as evolutionary
creationism, advocates that God guided the pro-
cess of evolution. It is a middle-ground between
Darwinism and religion. Supporters of theistic
predetermination believed that God created the
universe, as well as the natural laws within
it. This view is very similar to deistic evolution
(a position encompassing the belief that God cre-
ated the universe but has not interfered with it
since). Another form of theistic evolution pro-
posed that God was involved in the creation pro-
cess, either to start abiogenesis (spontaneous
generation of life from nonliving matter) or to
ensure that some mutations would have beneficial
Natural Selection
effects. Guided evolution posited that God used
evolution so that humans could evolve. This view,
similar to progressive creationism, comprised
belief that God extensively intervened in the pro-
cess of evolution, either by generating species or
by modifying gene sequences, and helping bene-
ficial mutations gain selective advantage. By the
beginning of the twentieth century, theistic evolu-
tion was rarely a part of scientific discussions, but
it kept a following among laymen.
Orthogenesis, or orthogenic (progressive,
straight-line) evolution, proposed that all organ-
isms carry an innate tendency to evolve towards a
predetermined, fixed goal. This teleological
hypothesis of a unilinear change of species
towards perfection was supported by many emi-
nent scientists who found proof for the argument
in the seemingly constant and gradual unidirec-
tional change within the fossil records. The term
was introduced in 1893 by Wilhelm Haacke, a
German zoologist. Orthogenesis was largely
discarded with the emergence of the modern syn-
thesis, but the idea that evolution represents pro-
gress remained popular. Although this is not true,
evolution does proceed in a seemingly linear way,
which is in accordance with the neo-Darwinism.
Vitalism proposed that all living organisms are
fundamentally different from the nonliving matter
due to containing a non-physical element
(sometimes referred to as the vital spark or
energy), or regulated by different principles
(Bechtel and Richardson 1998). The idea has
been discussed since ancient Egypt and ancient
Greece, through the Middle Ages to the nine-
teenth century. Among the supporters of this the-
ory was Louis Pasteur who, after conducting
several experiments, concluded that fermentation
was a vital action. By mid-twentieth century, the
majority of scientists had abandoned vitalism.
Saltationism or mutationism offered a faster
alternative to the Darwinian concept of gradual
change of natural selection, suggesting that new
species emerge as a result of large mutations. This
hypothesis was the basis of the mutation theory of
evolution. The central premise was that species go
through periods of fast mutation, possibly due to
environmental stress. These mutations could pro-
duce subspecies or new species in a single
9
generation. Modern science accepts the role of
mutation in evolution, albeit only as a supplier
of variation, which is then acted upon by natural
selection.
Catastrophism is a theory that hypothesizes
that the Earth was affected in the past by sudden
violent events (Turney and Brown 2007).
A French paleontologist Georges Cuvier pro-
posed that the various extinctions and the patterns
of faunal succession in the fossil record were
caused by these large-scale natural catastrophies.
He found evidence in the stratigraphic record that
such catastrophic events occurred but were sepa-
rated by long periods of stability. Modern biology
allows for catastrophism due to the evidence of a
large extinction event at the end of the Cretaceous
period, which led to a loss of approximately 70%
of all species (Renne et al. 2013).
Structuralism (biological or process structural-
ism) argued that evolution is not solely driven by
natural selection, but that important roles are
played by other mechanisms, which can in some
cases completely override natural selection. Struc-
turalism comprises several theories. Etienne
Geoffroy Saint-Hillarie’s law of compensation
suggests that all animals share an ideal pattern,
as evidenced by their homologous morphology.
This is not a result of evolution, but a product of
the law of nature. As certain parts become more
developed, others are reduced in compensation.
D’Arcy Wentworth Thompson’s universal laws of
form explain that the morphology of living organ-
isms often echoes inorganic structures. Adolf
Sellacher’s believed that fossils were the
so-called pneu structures and were determined
by mechanical inflation rather than natural selec-
tion. Gunter P. Wagner argued that homology and
biological novelty can be explained by a develop-
mental bias of structural constraints on embryonic
development. Stuart Kauffman proposed that self-
organization plays a role alongside natural selec-
tion in population dynamics, molecular evolution,
and morphogenesis. Michael Denton argued the
existence of the so-called Types – the basic forms
in nature, which were determined by the laws of
form. He believed that these universal, recurring
patterns were not shaped by natural selection but
by the self-organizing properties of matter and the
10
laws of nature. Stephen J. Gould and Richard
Lewontin suggested that many biological fea-
tures, which they compared to architectural span-
drels, do not always arise as direct results of
adaptation but can be consequences of evolution-
ary changes, or even exaptations. Brian Goodwin
proposed that natural patterns can be created by
spatial oscillations of chemical signals, similar to
the way in which morphogenetic fields operate
(Webster and Goodwin 1996). Extreme structur-
alists G.B. Muller and S.A. Newman held the
view that diversifications of species are dictated
by the physical laws of structure. They believed
that a relatively unconstrained, “pre-Mendelian”
phase in animal evolution existed in the early
stages of the multicellular life, suggesting possi-
ble explanations of how organisms evolved
genetic mechanisms (Newman et al. 2003).
The most popular alternative to Darwin’s the-
ory at the end of the nineteenth century was neo-
Lamarckism. In addition to Lamarck’s original
ideas, neo-Lamarckism included a proposition
that environment can directly alter organic struc-
tures. This was used as an explanation to why
organisms are adapted to their environments.
However, the largest flaw of this theory was the
complete lack of solid evidence suggesting the
inheritance of acquired traits. As the genetics
field progressively developed, neo-Lamarckism
began to lose support.
The Modern Synthesis (the
Neo-Darwinism)
The Modern Synthesis is the modern theory of
evolution that arose in the 1930s and 1940s from
the merger of Darwinian theory of evolution and
Mendelian genetics. It advocates the genetic basis
of evolution, describing it as a change in the allele
frequency within a population. The theory is also
based in the population genetics, and the work of
such as Ronald Aylmer Fisher, an English biolo-
gist and statistician, Sewall Wright, an American
geneticist, or John Burdon Sanderson Haldane, an
English-born Indian biologist and biostatistician.
Natural Selection
Central Contributors of the Modern Synthesis
Several distinguished scientists contributed to the
development of the Modern Synthesis.
Theodosius Grygorovych Dobzhansky, a
Ukrainian-American geneticist and evolutionary
biologist, greatly influenced the development of
the Modern Synthesis with his 1937 publication
Genetics and the Origin of Species. He proposed
that evolution is a change in the allele frequency
within a gene pool, explaining that natural selec-
tion works through mutation (Dobzhansky 1937).
Ernst Walter Mayr was an evolutionary biolo-
gist and a taxonomist. His main contribution to the
Modern Synthesis theory came from his
re-definition of the term species so as to include
a group of not only morphologically similar indi-
viduals but those who can breed only among
themselves. He also developed a theory of peri-
patric speciation – explaining the speciation that
arises from an isolated peripheral population
(Mayr 1954), which was the basis of the theory
of punctuated equilibrium, developed by Stephen
Jay Gould and Niles Eldredge (Eldredge and
Gould 1972). This theory proposed that most
changes that occurred in the evolutionary history
happened very rapidly, in short bursts of
branching speciation (i.e., cladogenesis) typically
lasting less than 100,000 years. After a species has
appeared in the fossil record, it will show very
little evolutionary change for the rest of its geo-
logical history, which they called stasis. The
opposite idea, proposing that evolution typically
occurs smoothly and uniformly, by the steady
gradual change of entire lineages is called phyletic
gradualism.
George Gaylord Simpson, an American pale-
ontologist, integrated paleontology with genetics
and natural selection. He developed methods of
analysis of the rates of evolution through time and
showed their variability. Furthermore, he intro-
duced the term of quantum evolution to explain
sudden emergences of lineages (Simpson 1944).
He suggested that microevolution of population
genetics offered an adequate explanation of the
macroevolution patterns observable by paleontol-
ogy. In his book Tempo and Mode in Evolution,
published in 1944, he defined tempo as the
Natural Selection
evolutionary acceleration and deceleration, the
evidence of which could both be found in the
fossil record. Simpson described mode as the pat-
tern of evolution, more encompassing than evolu-
tionary rates that were its basic factor. Simpson
noted that Gould’s and Eldredge’s punctuated
equilibrium theory is a reiteration of his idea of
quantum evolution.
George Ledyard Stebbins Jr. was an American
botanist and geneticist, and one of the leading
evolutionary biologists of the twentieth century.
In his work he combined genetics and Darwin’s
theory of evolution by natural selection to describ-
ing the speciation of plant species (Stebbins
1950). He was the first to discuss natural selection
in plants.
Julian Huxley, a British evolutionary biologist
and T.H. Huxley’s grandson, coined the terms
modern synthesis and evolutionary synthesis. In
his 1942 publication Evolution: The Modern Syn-
thesis, he integrated Darwin’s theory of evolution
with Mendelian genetics and population genetics.
He believed that different groups will differ in the
nature of their evolution, depending on the vari-
ous factors, for example their reproductive strate-
gies and genetic material (Huxley 1942). He
published numerous books in collaboration with
many of his eminent peers and worked on the
popularization of science and evolutionary biol-
ogy through many public appearances on televi-
sion and the radio.
Bernhard Rensch, a German evolutionary biol-
ogist, popularized the Modern Synthesis in Ger-
many. He studied environmental impacts on the
evolution of geographically isolated species.
Moreover, in his book Evolution above the spe-
cies level, published in 1947, he aimed to prove
that the major evolutionary trends and the forma-
tion of species and races are governed by the same
factors (Rensch 1960).
Differences Between the Modern Synthesis
and Darwin’s Theory
According to the premises of the Modern Synthe-
sis, natural selection remains a driving force of
evolution, but its power is joined with that of gene
flow (the transfer of alleles or genes between
populations, which happens as a result of
11
immigration or emigration of individuals), genetic
drift (the random change in allele frequency in a
population caused by random sampling), and
mutation (a change in the DNA sequence, which
can involve deletions, duplications, insertions, or
inversions (translocations)). The Modern Synthe-
sis provides an explanation of how genetic varia-
tion (i.e., variation in the alleles) persists. As
Darwin supported the theory of blending inheri-
tance, he struggled to explain this. Mendelian
inheritance gave a new perspective for tackling
this issue. Gregor Mendel was an Austrian scien-
tist and a monk who, after experimenting with pea
plants, showed that genes play a role in the traits
of an organism, and that those traits can be statis-
tically predicted using the knowledge of genetic
makeup of organism’s parents. According to
Mendel’s laws, alleles (i.e., DNA segments of
genes) do not merge and blend out. In reference
to traits, Mendel created the terms “dominant,”
describing the allele whose effect on organism’s
phenotype masks the contribution of the other
allele, and “recessive,” describing the allele
whose effect is masked by that of the other allele.
While Mendel’s work is the cornerstone of mod-
ern genetics, his views were not broadly accepted
or known until the beginning of the twentieth
century. Mendel was aware of Darwin’s work,
which he studied in great detail. There are a few
indications that Darwin could have been familiar
with Mendel’s work – portions of Mendel’s pub-
lications and references to them made by other
authors were included in books later found in
Darwin’s library and were thus likely available
to him. Mendel sent a copy of his 1865 paper
“Versuche uber Pflanzen-Hybriden” (“Experi-
ments in Plant Hybridization”) to 40 of the most
renowned nineteenth-century scientists, but it is
not known for certain whether or not Darwin was
one of the 40 scientists who received a copy, or
whether he read it.
According to Mendelian genetics, genetic var-
iation can occur both within populations and
among them. It allows the natural selection to
decrease or increase the frequency of alleles that
already exist. Genetic variation arises from the
formation of new alleles, alteration of the number
or position of genes, rapid reproduction, and
12
recombination between chromatids of homolo-
gous chromosomes during meiosis (a process
also known as the crossing over) in sexually
reproducing species. Some genetic variations
that arise can have a neutral effect, in which case
they will be neither selected for or against, and are
likely to remain in the population. In addition to
the variation within a population, species can also
exhibit geographic variation – variation in the
genotype of geographically separated
populations. These distinctions can arise either
due to a separation by environmental barriers or
because of the different selection pressures caused
by different environments. When a change in an
environment generates new available resources,
creates new challenges or opens new niches,
adaptive radiation (also known as divergent evo-
lution) can take place. It is a rapid diversification
of species into a multitude of new forms. One of
the most famous examples of an adaptive radia-
tion is the so-called Darwin’s finches. Occupying
fragmented landscape of the Galapagos islands,
they have diversified in their ecology and
morphology – most notably the size and shape
of their beaks, which provided Darwin with evi-
dence for his theory of evolution.
The Second Synthesis: Evolutionary
Developmental Biology
As molecular genetics continue to advance, a new
field of evolutionary developmental biology
started to emerge. The research of this field, infor-
mally referred to as “evo-devo,” aims to explain
evolution in terms of the genetic regulatory
network – a collection of molecular regulators
governing the gene expression of mRNA and pro-
teins, through the interaction with each other and
other cell substances. They have a central role in
the creation of body structures (morphogenesis).
The central premise of the field is the existence of
a subset of genes that control the organism’s
embryonic development, called the evo-devo
gene toolkit. These genes are conserved among
phyla and thought to date back to the last common
ancestor of all bilaterally symmetric animals
(animals that have mirror symmetry in the sagittal
Natural Selection
plane vertically dividing their bodies into left and
right halves, with each side comprising one of
each sense organ and limb pair). These genes
also tend to convergently (independently) evolve
the same function. According to evolutionary
developmental biology, variation does not arise
from mutation of gene sequences but from
mutation-driven changes in gene regulation. Var-
iation can thus result from a toolkit gene being
expressed in a different pattern (i.e., under-
expressed or non-expressed) or from it acquiring
a new function.
Darwin’s theory also left a mark on the devel-
opment of cell-biology and molecular-biology.
The so-called cellular Darwinism proposes that
stochasticity (or random variation) at the molecu-
lar level generates diversity in cell types, whereas
cell interactions import a characteristic order on
the developing embryo (Kupiec 2010). It is based
on the work of Wilhelm Roux, a German zoolo-
gist and embryologist, who believed that a Dar-
winian competition occurs at all levels of
organisms, from molecules to organs, during
embryonic development.
Social Darwinism
Darwin’s theory of evolution by natural selection
had implications for the social sciences. The term
social Darwinism refers to theories that apply his
concepts of natural evolution to human society.
Social Darwinism emerged in the late 1880s and
was first described by Oscar Schmidt, but the term
was used fairly rarely until 1944 when Richard
Hofstadter, an American historian, published a
book entitled Social Darwinism in American
Thought. Social evolution and cultural evolution
hypotheses were common during the Enlighten-
ment, as well as seventeenth-century thinkers, but
social Darwinism differs from the other theories
of social change as the ideas are drawn from
biology. In addition to believing that humans
also struggle for resources and that advantageous
physical and psychological traits are accumulated
over time, Darwin proposed that social instincts
also evolve through natural selection, which
Natural Selection
strengthens societies in which they occur (Darwin
1871).
Universal Darwinism
Universal or generalized Darwinism, also known
as the Darwinian metaphysics or the universal
selection theory, comprises a variety of extensions
of Darwin’s original theory of evolution, aiming
to explain evolution in an array of other disci-
plines. These extensions can be grouped
depending on whether they focus on the biologi-
cal implications of evolution (i.e., psychology or
medicine) or whether they discuss the evolution of
entities other than genes (i.e., computer science).
Gene-based extensions include evolutionary psy-
chology, evolutionary aesthetics, evolutionary
anthropology, evolutionary musicology, sociobi-
ology, human behavioral ecology, evolutionary
epistemology, evolutionary medicine, Darwinian
literary studies, Darwinian happiness, molecular
evolution, evolutionary linguistics, and other.
Examples of non-gene-based extensions include
quantum Darwinism, cosmological natural selec-
tion, memetics, cultural selection theory, evolu-
tionary economics, evolutionary ethics,
evolutionary art, evolutionary music, genetic
algorithms, dual inheritance theory, and neural
Darwinism.
Cross-References
▶ Adaptation
▶ Alfred Russel Wallace
▶ Catastrophism
▶ Charles Darwin
▶ Directional Selection
▶ Divergent Evolution
▶ Evolution
▶ Fisher
▶ Fitness
▶ Gene Flow
▶ Genetic Drift
▶ Genetic Variation
▶ Genotype
▶ Gregor Mendel
13
▶ Jean-Baptiste Lamarck
▶ Mendel’s Laws
▶ Mendelian Crosses
▶ Mutations
▶ Origin of Species
▶ Phenotype
▶ Population
▶ Quantitative Genetics
▶ Runaway Selection
▶ Sexual Selection
▶ Stabilizing Selection
▶ Structuralism
▶ Survival Value
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