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A book of readings from Wikipedia

Volume 1: An Introduction

Khalid Chraibi

04 septembre 2010
Part of the Biology series on Evolution
Mechanisms and processes
Genetic drift
Gene flow
Natural selection
Research and history
Evolutionary history of life
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact
Evolutionary biology fields
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Population genetics
Biology portal · v • d • e
Part of the Biology series on Genetics

Key components
History and topics
Classical genetics
Evolution · Molecular
Mendelian inheritance
Molecular genetics
DNA sequencing
Genetic engineering
Genomics · Topics
Medical genetics
Branches in genetics
Evolution Wikipedia the main articles
Introduction to evolution
Evolution as theory and fact
Evolutionary history of life
Timeline of evolution
History of evolutionary thought
On the Origin of Species
The Genetical Theory of Natural Selection
Modern evolutionary synthesis
Base concepts
Common descent
Evidence of common descent
Genetic drift
Gene flow
Natural selection
Molecular phylogenetics
Evolutionary developmental biology
Molecular evolution
Human evolution
Evolutionary psychology
Controversy and social impacts
Creation–evolution controversy
Objections to evolution
Intelligent design
Social effect of evolutionary theory
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Basic topics in evolutionary biology
Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary Canalisation · Modularity · Phenotypic plasticity

biology (Evo-devo)

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from ""

Index of evolutionary biology articles

From Wikipedia, the free encyclopedia

(Redirected from List of evolutionary biology topics)
Jump to: navigation, search
This is a list of topics in evolutionary biology.
Main article: Evolutionary biology
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z—
See also
This list is incomplete; you can help by expanding it.
[edit] A
abiogenesis - adaptation - adaptive radiation - allele - allele frequency - allopatric speciation -
altruism - anagenesis - Archaeopteryx - aquatic adaptation - artificial selection - atavism

[edit] B
biological organisation - Brassica oleracea - breed

[edit] C
Cambrian explosion - catagenesis - gene-centered view of evolution - cephalization - Sergei
Chetverikov - Chi square test - chronobiology - chronospecies - clade - cladistics - Climatic adaptation -
coalescent theory - co-evolution - Co-operation (evolution) - coefficient of relationship - common descent -
convergent evolution - creation-evolution controversy - cultivar - current research in evolutionary biology

[edit] D
Darwin (unit) - Charles Darwin - Darwinism - Darwin's finches - Richard Dawkins - directional
selection - Theodosius Dobzhansky - Dog breeding - Domestication - Domestication of the horse

[edit] E
ecological genetics - ecological selection - endosymbiosis - error threshold (evolution) - evidence of
common descent - evolution - evolutionary arms race - evolutionary capacitance - evolution of cetaceans -
evolution of complexity - evolution of the horse - evolution of mammalian auditory ossicles - evolution of
mammals - evolution of sex - evolution of sirenians - evolution of the eye - evolutionary developmental
biology - evolutionary neuroscience - evolutionary psychology - evolutionary radiation - evolutionary stable
strategy - evolutionary tree - evolvability - experimental evolution - exaptation - extinction

[edit] F
Joe Felsenstein - R.A. Fisher - Fisher's reproductive value - fitness - fitness landscape - E.B. Ford -

[edit] G
Galápagos Islands - gene - gene-centric view of evolution - gene duplication - gene flow - gene pool -
genetic drift - genetic hitchhiking - genetic recombination - genetic variation - genotype - genotype-
environment correlation - genotype-environment interaction - genotype-phenotype distinction - Stephen Jay
Gould - gradualism - Peter and Rosemary Grant - group selection

[edit] H
J. B. S. Haldane - W. D. Hamilton - Hardy-Weinberg principle - heredity - hierarchy of life - history of
evolutionary thought - horizontal gene transfer - human evolution - human evolutionary genetics -
homologous chromosomes - homology (biology) - Julian Huxley - human vestigiality
[edit] I
inclusive fitness - insect evolution - Invertebrate paleontology (a.k.a. invertebrate paleobiology or

[edit] K
kin selection - Motoo Kimura–Karyotype - Koinophilia

[edit] L
Jean-Baptiste Lamarck - Lamarckism - landrace - Language -Last common ancestor - Last universal
ancestor - Richard Lewontin - list of gene families - List of human evolution fossils - life-history theory - Wen-
Hsiung Li - living fossils - Charles Lyell

[edit] M
macroevolution - macromutation - The Major Transitions in Evolution - maladaptation - mass
extinctions - mating systems - John Maynard Smith - Ernst Mayr - Gregor Mendel - memetics - Mendelian
inheritance - microevolution - Micropaleontology (a.k.a. micropaleobiology) - Mitochondrial Eve - modern
evolutionary synthesis - molecular clock - molecular evolution - molecular phylogeny - molecular systematics
- Monogamy - most recent common ancestor - Hermann Joseph Muller - Muller's ratchet - mutation -
mutational meltdown
[edit] N
natural selection - Nature versus nurture - Neo-Darwinism - neutral theory of molecular evolution -
Baron Franz Nopcsa

[edit] O
Susumu Ohno - Aleksandr Oparin - On The Origin of Species - Origin of birds - orthologous genes

[edit] P
Paleoanthropology - Paleobiology - Paleobotany - Paleontology - Paleozoology - Paleozoology:
Vertebrates - Paleozoology: Invertebrates - parallel evolution - paraphyletic - particulate inheritance -
peppered moth - peppered moth evolution - peripatric speciation - phenotype - phylogenetics - phylogeny -
phylogenetic tree - pikaia - Plant evolution - polymorphism (biology) - population - Population bottleneck -
population dynamics - population genetics - preadaptation - prehistoric archaeology - Principles of Geology -
George R. Price - Price equation - punctuated equilibrium - Plant Evolutionary Developmental Biology

[edit] Q
quasispecies model

[edit] R
race (biology) - Red Queen - recapitulation theory - recombination - Bernhard Rensch
[edit] S
selection - selective breeding - Science of Evolution - sexual selection - sociobiology - species -
speciation - species flock - sperm competition - stabilizing selection - strain (biology) - subspecies - survival
of the fittest - Survival value - symbiogenesis - systematics - George Gaylord Simpson - G. Ledyard Stebbins

[edit] T
Tiktaalik - timeline of evolution - Trait (biological) - transgressive phenotype - transitional fossil -
transposon - Triangle of U

[edit] U
Uniformitarianism (science) - Unit of selection

[edit] V
variety (plant) - Vertebrate paleontology (a.k.a. vertebrate paleobiology or paleozoology) - viral
evolution - The Voyage of the Beagle - vestigiality

[edit] W
Alfred Russel Wallace - Wallace effect - Wallace Line - Wallacea - George C. Williams - Edward O.
Wilson - Sewall Wright
[edit] Y
Y-chromosomal Adam - Y-DNA haplogroups by ethnic groups

[edit] See also

• List of biology topics
• List of biochemistry topics
Retrieved from ""

Categories: Biology lists | Evolutionary biology | Indexes of articles

• This page was last modified on 21 August 2010 at 18:28.

Related topics
Evolutionary biology fields— Cladistics • Ecological genetics • Evolutionary development • Human
evolution • Molecular evolution • Phylogenetics • Population genetics
Mechanisms— Natural selection • Genetic drift • Gene flow
Outcomes— Adaptation • Co-evolution • Co-operation • Speciation • Extinction
Research and history— Evidence of common descent • Evolutionary history of life • History of evolutionary
thought • Modern evolutionary synthesis • Social effect of evolutionary theory • Objections to evolution

Evolutionary biology
Evolutionary biologists • Eugenics • Evolutionary biology literature • Evolutionary dynamics •
Evolutionary game theory • Extinction • History of evolutionary biology • Human evolution • Microbial
population biology • Molecular evolution • Mutation • Paleontology • Phylogenetics • Population genetics •
Evolutionary psychology • Selection • Sociobiology • Speciation • Symbiosis • Tree of life •

From Wikipedia, the free encyclopedia

Jump to: navigation, search

For other uses, see Universe (disambiguation).
Physical cosmology
Universe · Big Bang
Age of the universe
Timeline of the Big Bang
Ultimate fate of the universe
[show]Early universe

Inflation · Nucleosynthesis
GWB · Neutrino background
Cosmic microwave background

[show]Expanding universe

Redshift · Hubble's law

Metric expansion of space
Friedmann equations
FLRW metric
[show]Structure Formation

Shape of the Universe

Structure formation
Galaxy formation
Large-scale structure
Galaxy filaments


Lambda-CDM model
Dark energy · Dark matter


Timeline of cosmological theories

Future of an expanding universe


Observational cosmology
2dF · SDSS
COBE · BOOMERanG · WMAP · Planck

Isaac Newton · Einstein · Hawking · Friedman · Lemaître · Hubble · Penzias · Wilson · Gamow ·
Dicke · Zel'dovich · Mather · Rubin · Smoot· others
The universe is commonly defined as the totality of everything that exists,[1] including all physical
matter and energy, the planets, stars, galaxies, and the contents of intergalactic space,[2][3] although this
usage may differ with the context (see definitions, below). The term universe may be used in slightly different
contextual senses, denoting such concepts as the cosmos, the world, or nature.
Observations of earlier stages in the development of the universe, which can be seen at great
distances, suggest that the universe has been governed by the same physical laws and constants throughout
most of its extent and history.
• 1 History
• 2 Etymology, synonyms and definitions
• 2.1 Broadest definition: reality and probability
• 2.2 Definition as reality
• 2.3 Definition as connected space-time
• 2.4 Definition as observable reality
• 3 Size, age, contents, structure, and laws
• 3.1 Fine tuning
• 4 Historical models
• 4.1 Creation
• 4.2 Philosophical models
• 4.3 Astronomical models
• 5 Theoretical models
• 5.1 General theory of relativity
• 5.2 Special relativity and space-time
• 5.3 Solving Einstein's field equations
• 5.4 Big Bang model
• 6 Untestable proposals
• 6.1 Multiverse theory
[edit] History
Throughout recorded history, several cosmologies and cosmogonies have been proposed to account
for observations of the universe. The earliest quantitative geocentric models were developed by the ancient
Greeks, who proposed that the universe possesses infinite space and has existed eternally, but contains a
single set of concentric spheres of finite size – corresponding to the fixed stars, the Sun and various planets –
rotating about a spherical but unmoving Earth. Over the centuries, more precise observations and improved
theories of gravity led to Copernicus's heliocentric model and the Newtonian model of the Solar System,
respectively. Further improvements in astronomy led to the realization that the Solar System is embedded in
a galaxy composed of millions of stars, the Milky Way, and that other galaxies exist outside it, as far as
astronomical instruments can reach. Careful studies of the distribution of these galaxies and their spectral
lines have led to much of modern cosmology. Discovery of the red shift and cosmic microwave background
radiation revealed that the universe is expanding and apparently had a beginning.
This high-resolution image of the Hubble ultra deep field shows a diverse range of galaxies, each
consisting of billions of stars. The equivalent area of sky that the picture occupies is shown in the lower left
corner. The smallest, reddest galaxies, about 100, are some of the most distant galaxies to have been
imaged by an optical telescope, existing at the time shortly after the Big Bang.
According to the prevailing scientific model of the universe, known as the Big Bang, the universe
expanded from an extremely hot, dense phase called the Planck epoch, in which all the matter and energy of
the observable universe was concentrated. Since the Planck epoch, the universe has been expanding to its
present form, possibly with a brief period (less than 10 −32 seconds) of cosmic inflation. Several independent
experimental measurements support this theoretical expansion and, more generally, the Big Bang theory.
Recent observations indicate that this expansion is accelerating because of dark energy, and that most of the
matter in the universe may be in a form which cannot be detected by present instruments, and so is not
accounted for in the present models of the universe; this has been named dark matter. The imprecision of
current observations has hindered predictions of the ultimate fate of the universe.
Current interpretations of astronomical observations indicate that the age of the universe is 13.75
±0.17 billion years,[4] and that the diameter of the observable universe is at least 93 billion light years, or
8.80 × 1026 metres.[5] According to general relativity, space can expand faster than the speed of light,
although we can view only a small portion of the universe due to the limitation imposed by light speed. Since
we cannot observe space beyond the limitations of light (or any electromagnetic radiation), it is uncertain
whether the size of the universe is finite or infinite.

[edit] Etymology, synonyms and definitions

See also: Cosmos, Nature, World (philosophy), and Celestial spheres
The word universe derives from the Old French word Univers, which in turn derives from the Latin
word universum.[6] The Latin word was used by Cicero and later Latin authors in many of the same senses
as the modern English word is used.[7] The Latin word derives from the poetic contraction Unvorsum — first
used by Lucretius in Book IV (line 262) of his De rerum natura (On the Nature of Things) — which connects
un, uni (the combining form of unus', or "one") with vorsum, versum (a noun made from the perfect passive
participle of vertere, meaning "something rotated, rolled, changed").[7] Lucretius used the word in the sense
"everything rolled into one, everything combined into one".

Artistic rendition (highly exaggerated) of a Foucault pendulum showing that the Earth is not
stationary, but rotates.
An alternative interpretation of unvorsum is "everything rotated as one" or "everything rotated by
one". In this sense, it may be considered a translation of an earlier Greek word for the universe, περιφορά,
"something transported in a circle", originally used to describe a course of a meal, the food being carried
around the circle of dinner guests.[8] This Greek word refers to an early Greek model of the universe, in
which all matter was contained within rotating spheres centered on the Earth; according to Aristotle, the
rotation of the outermost sphere was responsible for the motion and change of everything within. It was
natural for the Greeks to assume that the Earth was stationary and that the heavens rotated about the Earth,
because careful astronomical and physical measurements (such as the Foucault pendulum) are required to
prove otherwise.
The most common term for "universe" among the ancient Greek philosophers from Pythagoras
onwards was τὸ πᾶν (The All), defined as all matter (τὸ ὅλον) and all space (τὸ κενόν).[9][10] Other
synonyms for the universe among the ancient Greek philosophers included κόσμος (meaning the world, the
cosmos) and φύσις (meaning Nature, from which we derive the word physics).[11] The same synonyms are
found in Latin authors (totum, mundus, natura)[12] and survive in modern languages, e.g., the German words
Das All, Weltall, and Natur for universe. The same synonyms are found in English, such as everything (as in
the theory of everything), the cosmos (as in cosmology), the world (as in the many-worlds hypothesis), and
Nature (as in natural laws or natural philosophy).[13]

[edit] Broadest definition: reality and probability

See also: Introduction to quantum mechanics, Interpretation of quantum mechanics, and Many-
worlds hypothesis
The broadest definition of the universe can be found in De divisione naturae by the medieval
philosopher and theologian Johannes Scotus Eriugena, who defined it as simply everything: everything that
is created and everything that is not created. Time is not considered in Eriugena's definition; thus, his
definition includes everything that exists, has existed and will exist, as well as everything that does not exist,
has never existed and will never exist. This all-embracing definition was not adopted by most later
philosophers, but something not entirely dissimilar reappears in quantum physics, perhaps most obviously in
the path-integral formulation of Feynman.[14] According to that formulation, the probability amplitudes for the
various outcomes of an experiment given a perfectly defined initial state of the system are determined by
summing over all possible paths by which the system could progress from the initial to final state. Naturally,
an experiment can have only one outcome; in other words, only one possible outcome is made real in this
universe, via the mysterious process of quantum measurement, also known as the collapse of the
wavefunction (but see the many-worlds hypothesis below in the Multiverse section). In this well-defined
mathematical sense, even that which does not exist (all possible paths) can influence that which does finally
exist (the experimental measurement). As a specific example, every electron is intrinsically identical to every
other; therefore, probability amplitudes must be computed allowing for the possibility that they exchange
positions, something known as exchange symmetry. This conception of the universe embracing both the
existent and the non-existent loosely parallels the Buddhist doctrines of shunyata and interdependent
development of reality, and Gottfried Leibniz's more modern concepts of contingency and the identity of

[edit] Definition as reality

See also: Reality and Physics
More customarily, the universe is defined as everything that exists, has existed, and will exist[ citation
needed]. According to this definition and our present understanding, the universe consists of three elements:
space and time, collectively known as space-time or the vacuum; matter and various forms of energy and
momentum occupying space-time; and the physical laws that govern the first two. These elements will be
discussed in greater detail below. A related definition of the term universe is everything that exists at a single
moment of cosmological time, such as the present, as in the sentence "The universe is now bathed uniformly
in microwave radiation".
The three elements of the universe (spacetime, matter-energy, and physical law) correspond roughly
to the ideas of Aristotle. In his book The Physics (Φυσικῆς, from which we derive the word "physics"),
Aristotle divided τὸ πᾶν (everything) into three roughly analogous elements: matter (the stuff of which the
universe is made), form (the arrangement of that matter in space) and change (how matter is created,
destroyed or altered in its properties, and similarly, how form is altered). Physical laws were conceived as the
rules governing the properties of matter, form and their changes. Later philosophers such as Lucretius,
Averroes, Avicenna and Baruch Spinoza altered or refined these divisions[citation needed]; for example,
Averroes and Spinoza discern natura naturans (the active principles governing the universe) from natura
naturata, the passive elements upon which the former act.

[edit] Definition as connected space-time

See also: Chaotic Inflation theory
It is possible to conceive of disconnected space-times, each existing but unable to interact with one
another. An easily visualized metaphor is a group of separate soap bubbles, in which observers living on one
soap bubble cannot interact with those on other soap bubbles, even in principle. According to one common
terminology, each "soap bubble" of space-time is denoted as a universe, whereas our particular space-time
is denoted as the universe, just as we call our moon the Moon. The entire collection of these separate space-
times is denoted as the multiverse.[15] In principle, the other unconnected universes may have different
dimensionalities and topologies of space-time, different forms of matter and energy, and different physical
laws and physical constants, although such possibilities are currently speculative.

[edit] Definition as observable reality

See also: Observable universe and Observational cosmology
According to a still-more-restrictive definition, the universe is everything within our connected space-
time that could have a chance to interact with us and vice versa.[ citation needed] According to the general
theory of relativity, some regions of space may never interact with ours even in the lifetime of the universe,
due to the finite speed of light and the ongoing expansion of space. For example, radio messages sent from
Earth may never reach some regions of space, even if the universe would live forever; space may expand
faster than light can traverse it. It is worth emphasizing that those distant regions of space are taken to exist
and be part of reality as much as we are; yet we can never interact with them. The spatial region within which
we can affect and be affected is denoted as the observable universe. Strictly speaking, the observable
universe depends on the location of the observer. By traveling, an observer can come into contact with a
greater region of space-time than an observer who remains still, so that the observable universe for the
former is larger than for the latter. Nevertheless, even the most rapid traveler may not be able to interact with
all of space. Typically, the observable universe is taken to mean the universe observable from our vantage
point in the Milky Way Galaxy.

[edit] Size, age, contents, structure, and laws

Main articles: Observable universe, Age of the universe, Large-scale structure of the universe, and
Abundance of the chemical elements
The universe is very large and possibly infinite in volume. The region visible from Earth (the
observable universe) is about 92 billion light years across,[16] based on where the expansion of space has
taken the most distant objects observed. For comparison, the diameter of a typical galaxy is only 30,000
light-years, and the typical distance between two neighboring galaxies is only 3 million light-years.[17] As an
example, our Milky Way Galaxy is roughly 100,000 light years in diameter,[18] and our nearest sister galaxy,
the Andromeda Galaxy, is located roughly 2.5 million light years away.[19] There are probably more than 100
billion (1011) galaxies in the observable universe.[20] Typical galaxies range from dwarfs with as few as ten
million[21] (107) stars up to giants with one trillion[22] (1012) stars, all orbiting the galaxy's center of mass.
Thus, a very rough estimate from these numbers would suggest there are around one sextillion (1021) stars
in the observable universe; though a 2003 study by Australian National University astronomers resulted in a
figure of 70 sextillion (7 x 1022)[23].
The universe is believed to be mostly composed of dark energy and dark matter, both of which are
poorly understood at present. Less than 5% of the universe is ordinary matter, a relatively small perturbation.
The observable matter is spread uniformly (homogeneously) throughout the universe, when
averaged over distances longer than 300 million light-years.[24] However, on smaller length-scales, matter is
observed to form "clumps", i.e., to cluster hierarchically; many atoms are condensed into stars, most stars
into galaxies, most galaxies into clusters, superclusters and, finally, the largest-scale structures such as the
Great Wall of galaxies. The observable matter of the universe is also spread isotropically, meaning that no
direction of observation seems different from any other; each region of the sky has roughly the same content.
[25] The universe is also bathed in a highly isotropic microwave radiation that corresponds to a thermal
equilibrium blackbody spectrum of roughly 2.725 kelvin.[26] The hypothesis that the large-scale universe is
homogeneous and isotropic is known as the cosmological principle,[27] which is supported by astronomical
The present overall density of the universe is very low, roughly 9.9 × 10−30 grams per cubic
centimetre. This mass-energy appears to consist of 73% dark energy, 23% cold dark matter and 4% ordinary
matter. Thus the density of atoms is on the order of a single hydrogen atom for every four cubic meters of
volume.[28] The properties of dark energy and dark matter are largely unknown. Dark matter gravitates as
ordinary matter, and thus works to slow the expansion of the universe; by contrast, dark energy accelerates
its expansion.
The universe is old and evolving. The most precise estimate of the universe's age is 13.73±0.12
billion years old, based on observations of the cosmic microwave background radiation.[29] Independent
estimates (based on measurements such as radioactive dating) agree, although they are less precise,
ranging from 11–20 billion years[30] to 13–15 billion years.[31] The universe has not been the same at all
times in its history; for example, the relative populations of quasars and galaxies have changed and space
itself appears to have expanded. This expansion accounts for how Earth-bound scientists can observe the
light from a galaxy 30 billion light years away, even if that light has traveled for only 13 billion years; the very
space between them has expanded. This expansion is consistent with the observation that the light from
distant galaxies has been redshifted; the photons emitted have been stretched to longer wavelengths and
lower frequency during their journey. The rate of this spatial expansion is accelerating, based on studies of
Type Ia supernovae and corroborated by other data.
The relative fractions of different chemical elements — particularly the lightest atoms such as
hydrogen, deuterium and helium — seem to be identical throughout the universe and throughout its
observable history.[32] The universe seems to have much more matter than antimatter, an asymmetry
possibly related to the observations of CP violation.[33] The universe appears to have no net electric charge,
and therefore gravity appears to be the dominant interaction on cosmological length scales. The universe
also appears to have neither net momentum nor angular momentum. The absence of net charge and
momentum would follow from accepted physical laws (Gauss's law and the non-divergence of the stress-
energy-momentum pseudotensor, respectively), if the universe were finite.[34]

The elementary particles from which the universe is constructed. Six leptons and six quarks comprise
most of the matter; for example, the protons and neutrons of atomic nuclei are composed of quarks, and the
ubiquitous electron is a lepton. These particles interact via the gauge bosons shown in the middle row, each
corresponding to a particular type of gauge symmetry. The Higgs boson (as yet unobserved) is believed to
confer mass on the particles with which it is connected. The graviton, a supposed gauge boson for gravity, is
not shown.
The universe appears to have a smooth space-time continuum consisting of three spatial dimensions
and one temporal (time) dimension. On the average, space is observed to be very nearly flat (close to zero
curvature), meaning that Euclidean geometry is experimentally true with high accuracy throughout most of
the Universe.[35] Spacetime also appears to have a simply connected topology, at least on the length-scale
of the observable universe. However, present observations cannot exclude the possibilities that the universe
has more dimensions and that its spacetime may have a multiply connected global topology, in analogy with
the cylindrical or toroidal topologies of two-dimensional spaces.[36]
The universe appears to behave in a manner that regularly follows a set of physical laws and
physical constants.[37] According to the prevailing Standard Model of physics, all matter is composed of
three generations of leptons and quarks, both of which are fermions. These elementary particles interact via
at most three fundamental interactions: the electroweak interaction which includes electromagnetism and the
weak nuclear force; the strong nuclear force described by quantum chromodynamics; and gravity, which is
best described at present by general relativity. The first two interactions can be described by renormalized
quantum field theory, and are mediated by gauge bosons that correspond to a particular type of gauge
symmetry. A renormalized quantum field theory of general relativity has not yet been achieved, although
various forms of string theory seem promising. The theory of special relativity is believed to hold throughout
the universe, provided that the spatial and temporal length scales are sufficiently short; otherwise, the more
general theory of general relativity must be applied. There is no explanation for the particular values that
physical constants appear to have throughout our universe, such as Planck's constant h or the gravitational
constant G. Several conservation laws have been identified, such as the conservation of charge, momentum,
angular momentum and energy; in many cases, these conservation laws can be related to symmetries or
mathematical identities.

[edit] Fine tuning

Main article: fine-tuned universe
It appears that many of properties of the universe have special values in the sense that a universe
where these properties only differ slightly would not be able to support intelligent life.[38][39] Not all scientists
agree that this fine-tuning exists.[40][41] In particular, it is not known under what conditions intelligent life
could form and what form or shape that would take. A relevant observation in this discussion is that existence
of an observer to observe fine-tuning, requires that the universe supports intelligent life. As such the
conditional probability of observing a universe that is fine-tuned to support intelligent life is 1. This
observation is known as the anthropic principle and is particularly relevant if the creation of the universe was
probabilistic or if multiple universes with a variety of properties exist (see below).

[edit] Historical models

See also: Cosmology and Timeline of cosmology
Many models of the cosmos (cosmologies) and its origin (cosmogonies) have been proposed, based
on the then-available data and conceptions of the universe. Historically, cosmologies and cosmogonies were
based on narratives of gods acting in various ways. Theories of an impersonal universe governed by physical
laws were first proposed by the Greeks and Indians. Over the centuries, improvements in astronomical
observations and theories of motion and gravitation led to ever more accurate descriptions of the universe.
The modern era of cosmology began with Albert Einstein's 1915 general theory of relativity, which made it
possible to quantitatively predict the origin, evolution, and conclusion of the universe as a whole. Most
modern, accepted theories of cosmology are based on general relativity and, more specifically, the predicted
Big Bang; however, still more careful measurements are required to determine which theory is correct.

[edit] Creation
Main articles: Creation myth and Creator deity

Sumerian account of the creatrix goddess Nammu, the precursor of the Assyrian goddess Tiamat;
perhaps the earliest surviving creation myth.
Many cultures have stories describing the origin of the world, which may be roughly grouped into
common types. In one type of story, the world is born from a world egg; such stories include the Finnish epic
poem Kalevala, the Chinese story of Pangu or the Indian Brahmanda Purana. In related stories, the creation
idea is caused by a single entity emanating or producing something by his or herself, as in the Tibetan
Buddhism concept of Adi-Buddha, the ancient Greek story of Gaia (Mother Earth), the Aztec goddess
Coatlicue myth, the ancient Egyptian god Atum story, or the Genesis creation narrative. In another type of
story, the world is created from the union of male and female deities, as in the Maori story of Rangi and
Papa. In other stories, the universe is created by crafting it from pre-existing materials, such as the corpse of
a dead god — as from Tiamat in the Babylonian epic Enuma Elish or from the giant Ymir in Norse mythology –
or from chaotic materials, as in Izanagi and Izanami in Japanese mythology. In other stories, the universe
emanates from fundamental principles, such as Brahman and Prakrti, or the yin and yang of the Tao.

[edit] Philosophical models

Further information: Cosmology
See also: Pre-Socratic philosophy, Physics (Aristotle), Hindu cosmology, Islamic cosmology, and
From the 6th century BCE, the pre-Socratic Greek philosophers developed the earliest known
philosophical models of the universe. The earliest Greek philosophers noted that appearances can be
deceiving, and sought to understand the underlying reality behind the appearances. In particular, they noted
the ability of matter to change forms (e.g., ice to water to steam) and several philosophers proposed that all
the apparently different materials of the world (wood, metal, etc.) are all different forms of a single material,
the arche. The first to do so was Thales, who called this material Water. Following him, Anaximenes called it
Air, and posited that there must be attractive and repulsive forces that cause the arche to condense or
dissociate into different forms. Empedocles proposed that multiple fundamental materials were necessary to
explain the diversity of the universe, and proposed that all four classical elements (Earth, Air, Fire and Water)
existed, albeit in different combinations and forms. This four-element theory was adopted by many of the
subsequent philosophers. Some philosophers before Empedocles advocated less material things for the
arche; Heraclitus argued for a Logos, Pythagoras believed that all things were composed of numbers,
whereas Thales' student, Anaximander, proposed that everything was composed of a chaotic substance
known as apeiron, roughly corresponding to the modern concept of a quantum foam. Various modifications of
the apeiron theory were proposed, most notably that of Anaxagoras, which proposed that the various matter
in the world was spun off from a rapidly rotating apeiron, set in motion by the principle of Nous (Mind). Still
other philosophers — most notably Leucippus and Democritus — proposed that the universe was composed of
indivisible atoms moving through empty space, a vacuum; Aristotle opposed this view ("Nature abhors a
vacuum") on the grounds that resistance to motion increases with density; hence, empty space should offer
no resistance to motion, leading to the possibility of infinite speed.
Although Heraclitus argued for eternal change, his quasi-contemporary Parmenides made the radical
suggestion that all change is an illusion, that the true underlying reality is eternally unchanging and of a
single nature. Parmenides denoted this reality as τὸ ἐν (The One). Parmenides' theory seemed implausible
to many Greeks, but his student Zeno of Elea challenged them with several famous paradoxes. Aristotle
resolved these paradoxes by developing the notion of an infinitely divisible continuum, and applying it to
space and time.
The Indian philosopher Kanada, founder of the Vaisheshika school, developed a theory of atomism
and proposed that light and heat were varieties of the same substance.[42] In the 5th century AD, the
Buddhist atomist philosopher Dignāga proposed atoms to be point-sized, durationless, and made of energy.
They denied the existence of substantial matter and proposed that movement consisted of momentary
flashes of a stream of energy.[43]
The theory of temporal finitism was inspired by the doctrine of creation shared by the three
Abrahamic religions: Judaism, Christianity and Islam. The Christian philosopher, John Philoponus, presented
the philosophical arguments against the ancient Greek notion of an infinite past. Philoponus' arguments
against an infinite past were used by the early Muslim philosopher, Al-Kindi (Alkindus); the Jewish
philosopher, Saadia Gaon (Saadia ben Joseph); and the Muslim theologian, Al-Ghazali (Algazel). They
employed two logical arguments against an infinite past, the first being the "argument from the impossibility
of the existence of an actual infinite", which states:[44]
"An actual infinite cannot exist."
"An infinite temporal regress of events is an actual infinite."

" An infinite temporal regress of events cannot exist."

The second argument, the "argument from the impossibility of completing an actual infinite by
successive addition", states:[44]
"An actual infinite cannot be completed by successive addition."
"The temporal series of past events has been completed by successive addition."

" The temporal series of past events cannot be an actual infinite."

Both arguments were adopted by later Christian philosophers and theologians, and the second
argument in particular became more famous after it was adopted by Immanuel Kant in his thesis of the first
antinomy concerning time.[44]

[edit] Astronomical models

Main article: History of astronomy
Astronomical models of the universe were proposed soon after astronomy began with the Babylonian
astronomers, who viewed the universe as a flat disk floating in the ocean, and this forms the premise for early
Greek maps like those of Anaximander and Hecataeus of Miletus.
Later Greek philosophers, observing the motions of the heavenly bodies, were concerned with
developing models of the universe based more profoundly on empirical evidence. The first coherent model
was proposed by Eudoxus of Cnidos. According to this model, space and time are infinite and eternal, the
Earth is spherical and stationary, and all other matter is confined to rotating concentric spheres. This model
was refined by Callippus and Aristotle, and brought into nearly perfect agreement with astronomical
observations by Ptolemy. The success of this model is largely due to the mathematical fact that any function
(such as the position of a planet) can be decomposed into a set of circular functions (the Fourier modes).
However, not all Greek scientists accepted the geocentric model of the universe. The Pythagorean
philosopher Philolaus postulated that at the center of the universe was a "central fire" around which the
Earth, Sun, Moon and Planets revolved in uniform circular motion.[45] The Greek astronomer Aristarchus of
Samos was the first known individual to propose a heliocentric model of the universe. Though the original
text has been lost, a reference in Archimedes' book The Sand Reckoner describes Aristarchus' heliocentric
theory. Archimedes wrote: (translated into English)
You King Gelon are aware the 'universe' is the name given by most astronomers to the sphere
the center of which is the center of the Earth, while its radius is equal to the straight line between
the center of the Sun and the center of the Earth. This is the common account as you have
heard from astronomers. But Aristarchus has brought out a book consisting of certain
hypotheses, wherein it appears, as a consequence of the assumptions made, that the universe
is many times greater than the 'universe' just mentioned. His hypotheses are that the fixed stars
and the Sun remain unmoved, that the Earth revolves about the Sun on the circumference of a
circle, the Sun lying in the middle of the orbit, and that the sphere of fixed stars, situated about
the same center as the Sun, is so great that the circle in which he supposes the Earth to revolve
bears such a proportion to the distance of the fixed stars as the center of the sphere bears to its

Aristarchus thus believed the stars to be very far away, and saw this as the reason why there was no
visible parallax, that is, an observed movement of the stars relative to each other as the Earth moved around
the Sun. The stars are in fact much farther away than the distance that was generally assumed in ancient
times, which is why stellar parallax is only detectable with telescopes. The geocentric model, consistent with
planetary parallax, was assumed to be an explanation for the unobservability of the parallel phenomenon,
stellar parallax. The rejection of the heliocentric view was apparently quite strong, as the following passage
from Plutarch suggests (On the Apparent Face in the Orb of the Moon):
Cleanthes [a contemporary of Aristarchus and head of the Stoics] thought it was the duty of the
Greeks to indict Aristarchus of Samos on the charge of impiety for putting in motion the Hearth
of the universe [i.e. the earth], . . . supposing the heaven to remain at rest and the earth to
revolve in an oblique circle, while it rotates, at the same time, about its own axis. [1]
The only other astronomer from antiquity known by name who supported Aristarchus' heliocentric
model was Seleucus of Seleucia, a Hellenized Babylonian astronomer who lived a century after Aristarchus.
[46][47][48] According to Plutarch, Seleucus was the first to prove the heliocentric system through reasoning,
but it is not known what arguments he used. Seleucus' arguments for a heliocentric theory were probably
related to the phenomenon of tides.[49] According to Strabo (1.1.9), Seleucus was the first to state that the
tides are due to the attraction of the Moon, and that the height of the tides depends on the Moon's position
relative to the Sun.[50] Alternatively, he may have proved the heliocentric theory by determining the
constants of a geometric model for the heliocentric theory and by developing methods to compute planetary
positions using this model, like what Nicolaus Copernicus later did in the 16th century.[51] During the Middle
Ages, heliocentric models may have also been proposed by the Indian astronomer, Aryabhata,[52] and by
the Persian astronomers, Albumasar[53] and Al-Sijzi.[54]
Model of the Copernican universe by Thomas Digges in 1576, with the amendment that the stars are
no longer confined to a sphere, but spread uniformly throughout the space surrounding the planets.
The Aristotelian model was accepted in the Western world for roughly two millennia, until Copernicus
revived Aristarchus' theory that the astronomical data could be explained more plausibly if the earth rotated
on its axis and if the sun were placed at the center of the universe.
“ In the center rests the sun. For who would place this lamp of a very beautiful ”
temple in another or better place than this wherefrom it can illuminate everything at the
same time?

—Copernicus, in Chapter 10, Book 1 of De Revolutionibus Orbium Coelestrum (1543)

As noted by Copernicus himself, the suggestion that the Earth rotates was very old, dating at least to
Philolaus (c. 450 BC), Heraclides Ponticus (c. 350 BC) and Ecphantus the Pythagorean. Roughly a century
before Copernicus, Christian scholar Nicholas of Cusa also proposed that the Earth rotates on its axis in his
book, On Learned Ignorance (1440).[55] Aryabhata (476–550), Brahmagupta (598–668), Albumasar and Al-
Sijzi, also proposed that the Earth rotates on its axis.[ citation needed] The first empirical evidence for the
Earth's rotation on its axis, using the phenomenon of comets, was given by Tusi (1201–1274) and Ali Kuşçu
(1403–1474).[citation needed] Tusi, however, continued to support the Aristotelian universe, thus Kuşçu was
the first to refute the Aristotelian notion of a stationary Earth on an empirical basis, similar to how Copernicus
later justified the Earth's rotation. Al-Birjandi (d. 1528) further developed a theory of "circular inertia" to
explain the Earth's rotation, similar to how Galileo Galilei explained it.[56][57]
Johannes Kepler published the Rudolphine Tables containing a star catalog and planetary tables
using Tycho Brahe's measurements.
Copernicus' heliocentric model allowed the stars to be placed uniformly through the (infinite) space
surrounding the planets, as first proposed by Thomas Digges in his Perfit Description of the Caelestiall Orbes
according to the most aunciente doctrine of the Pythagoreans, latelye revived by Copernicus and by
Geometricall Demonstrations approved (1576).[58] Giordano Bruno accepted the idea that space was infinite
and filled with solar systems similar to our own; for the publication of this view, he was burned at the stake in
the Campo dei Fiori in Rome on 17 February 1600.[58]
This cosmology was accepted provisionally by Isaac Newton, Christiaan Huygens and later
scientists,[58] although it had several paradoxes that were resolved only with the development of general
relativity. The first of these was that it assumed that space and time were infinite, and that the stars in the
universe had been burning forever; however, since stars are constantly radiating energy, a finite star seems
inconsistent with the radiation of infinite energy. Secondly, Edmund Halley (1720)[59] and Jean-Philippe de
Cheseaux (1744)[60] noted independently that the assumption of an infinite space filled uniformly with stars
would lead to the prediction that the nighttime sky would be as bright as the sun itself; this became known as
Olbers' paradox in the 19th century.[61] Third, Newton himself showed that an infinite space uniformly filled
with matter would cause infinite forces and instabilities causing the matter to be crushed inwards under its
own gravity.[58] This instability was clarified in 1902 by the Jeans instability criterion.[62] One solution to
these latter two paradoxes is the Charlier universe, in which the matter is arranged hierarchically (systems of
orbiting bodies that are themselves orbiting in a larger system, ad infinitum) in a fractal way such that the
universe has a negligibly small overall density; such a cosmological model had also been proposed earlier in
1761 by Johann Heinrich Lambert.[63] A significant astronomical advance of the 18th century was the
realization by Thomas Wright, Immanuel Kant and others that stars are not distributed uniformly throughout
space; rather, they are grouped into galaxies.[64]
The modern era of physical cosmology began in 1917, when Albert Einstein first applied his general
theory of relativity to model the structure and dynamics of the universe.[65] This theory and its implications
will be discussed in more detail in the following section.
[edit] Theoretical models
High-precision test of general relativity by the Cassini space probe (artist's impression): radio signals
sent between the Earth and the probe (green wave) are delayed by the warping of space and time (blue
lines) due to the Sun's mass.
Of the four fundamental interactions, gravitation is dominant at cosmological length scales; that is,
the other three forces are believed to play a negligible role in determining structures at the level of planets,
stars, galaxies and larger-scale structures. Since all matter and energy gravitate, gravity's effects are
cumulative; by contrast, the effects of positive and negative charges tend to cancel one another, making
electromagnetism relatively insignificant on cosmological length scales. The remaining two interactions, the
weak and strong nuclear forces, decline very rapidly with distance; their effects are confined mainly to sub-
atomic length scales.

[edit] General theory of relativity

Main articles: Introduction to general relativity, General relativity, and Einstein's field equations
Given gravitation's predominance in shaping cosmological structures, accurate predictions of the
universe's past and future require an accurate theory of gravitation. The best theory available is Albert
Einstein's general theory of relativity, which has passed all experimental tests hitherto. However, since
rigorous experiments have not been carried out on cosmological length scales, general relativity could
conceivably be inaccurate. Nevertheless, its cosmological predictions appear to be consistent with
observations, so there is no compelling reason to adopt another theory.
General relativity provides of a set of ten nonlinear partial differential equations for the spacetime
metric (Einstein's field equations) that must be solved from the distribution of mass-energy and momentum
throughout the universe. Since these are unknown in exact detail, cosmological models have been based on
the cosmological principle, which states that the universe is homogeneous and isotropic. In effect, this
principle asserts that the gravitational effects of the various galaxies making up the universe are equivalent to
those of a fine dust distributed uniformly throughout the universe with the same average density. The
assumption of a uniform dust makes it easy to solve Einstein's field equations and predict the past and future
of the universe on cosmological time scales.
Einstein's field equations include a cosmological constant (Λ),[65][66] that corresponds to an energy
density of empty space.[67] Depending on its sign, the cosmological constant can either slow (negative Λ) or
accelerate (positive Λ) the expansion of the universe. Although many scientists, including Einstein, had
speculated that Λ was zero,[68] recent astronomical observations of type Ia supernovae have detected a
large amount of "dark energy" that is accelerating the universe's expansion.[69] Preliminary studies suggest
that this dark energy corresponds to a positive Λ, although alternative theories cannot be ruled out as yet.[70]
Russian physicist Zel'dovich suggested that Λ is a measure of the zero-point energy associated with virtual
particles of quantum field theory, a pervasive vacuum energy that exists everywhere, even in empty space.
[71] Evidence for such zero-point energy is observed in the Casimir effect.

[edit] Special relativity and space-time

Main articles: Introduction to special relativity and Special relativity
Only its length L is intrinsic to the rod (shown in black); coordinate differences between its endpoints
(such as Δx, Δy or Δξ, Δη) depend on their frame of reference (depicted in blue and red, respectively).
The universe has at least three spatial and one temporal (time) dimension. It was long thought that
the spatial and temporal dimensions were different in nature and independent of one another. However,
according to the special theory of relativity, spatial and temporal separations are interconvertible (within
limits) by changing one's motion.
To understand this interconversion, it is helpful to consider the analogous interconversion of spatial
separations along the three spatial dimensions. Consider the two endpoints of a rod of length L. The length
can be determined from the differences in the three coordinates Δx, Δy and Δz of the two endpoints in a
given reference frame
L2 = Δx2 + Δy2 + Δz2
using the Pythagorean theorem. In a rotated reference frame, the coordinate differences differ, but
they give the same length
L2 = Δξ2 + Δη2 + Δζ2.
Thus, the coordinates differences (Δx, Δy, Δz) and (Δξ, Δη, Δζ) are not intrinsic to the rod, but merely
reflect the reference frame used to describe it; by contrast, the length L is an intrinsic property of the rod. The
coordinate differences can be changed without affecting the rod, by rotating one's reference frame.
The analogy in spacetime is called the interval between two events; an event is defined as a point in
spacetime, a specific position in space and a specific moment in time. The spacetime interval between two
events is given by
where c is the speed of light. According to special relativity, one can change a spatial and time
separation (L1, Δt1) into another (L2, Δt2) by changing one's reference frame, as long as the change
maintains the spacetime interval s. Such a change in reference frame corresponds to changing one's motion;
in a moving frame, lengths and times are different from their counterparts in a stationary reference frame.
The precise manner in which the coordinate and time differences change with motion is described by the
Lorentz transformation.

[edit] Solving Einstein's field equations

See also: Big Bang and Ultimate fate of the universe
The distances between the spinning galaxies increase with time, but the distances between the stars
within each galaxy stay roughly the same, due to their gravitational interactions. This animation illustrates a
closed Friedmann universe with zero cosmological constant Λ; such a universe oscillates between a Big
Bang and a Big Crunch.
Animation illustrating the metric expansion of the universe
In non-Cartesian (non-square) or curved coordinate systems, the Pythagorean theorem holds only on
infinitesimal length scales and must be augmented with a more general metric tensor gμν, which can vary
from place to place and which describes the local geometry in the particular coordinate system. However,
assuming the cosmological principle that the universe is homogeneous and isotropic everywhere, every point
in space is like every other point; hence, the metric tensor must be the same everywhere. That leads to a
single form for the metric tensor, called the Friedmann-Lemaître-Robertson-Walker metric

where (r, θ, φ) correspond to a spherical coordinate system. This metric has only two undetermined
parameters: an overall length scale R that can vary with time, and a curvature index k that can be only 0, 1 or
−1, corresponding to flat Euclidean geometry, or spaces of positive or negative curvature. In cosmology,
solving for the history of the universe is done by calculating R as a function of time, given k and the value of
the cosmological constant Λ, which is a (small) parameter in Einstein's field equations. The equation
describing how R varies with time is known as the Friedmann equation, after its inventor, Alexander
The solutions for R(t) depend on k and Λ, but some qualitative features of such solutions are general.
First and most importantly, the length scale R of the universe can remain constant only if the universe is
perfectly isotropic with positive curvature (k=1) and has one precise value of density everywhere, as first
noted by Albert Einstein. However, this equilibrium is unstable and since the universe is known to be
inhomogeneous on smaller scales, R must change, according to general relativity. When R changes, all the
spatial distances in the universe change in tandem; there is an overall expansion or contraction of space
itself. This accounts for the observation that galaxies appear to be flying apart; the space between them is
stretching. The stretching of space also accounts for the apparent paradox that two galaxies can be 40 billion
light years apart, although they started from the same point 13.7 billion years ago and never moved faster
than the speed of light.
Second, all solutions suggest that there was a gravitational singularity in the past, when R goes to
zero and matter and energy became infinitely dense. It may seem that this conclusion is uncertain since it is
based on the questionable assumptions of perfect homogeneity and isotropy (the cosmological principle) and
that only the gravitational interaction is significant. However, the Penrose-Hawking singularity theorems show
that a singularity should exist for very general conditions. Hence, according to Einstein's field equations, R
grew rapidly from an unimaginably hot, dense state that existed immediately following this singularity (when
R had a small, finite value); this is the essence of the Big Bang model of the universe. A common
misconception is that the Big Bang model predicts that matter and energy exploded from a single point in
space and time; that is false. Rather, space itself was created in the Big Bang and imbued with a fixed
amount of energy and matter distributed uniformly throughout; as space expands (i.e., as R(t) increases), the
density of that matter and energy decreases.
Space has no boundary – that is empirically more certain than any external observation. However,
that does not imply that space is infinite...(translated, original German)

Bernhard Riemann (Habilitationsvortrag, 1854)

Third, the curvature index k determines the sign of the mean spatial curvature of spacetime averaged
over length scales greater than a billion light years. If k=1, the curvature is positive and the universe has a
finite volume. Such universes are often visualized as a three-dimensional sphere S3 embedded in a four-
dimensional space. Conversely, if k is zero or negative, the universe may have infinite volume, depending on
its overall topology. It may seem counter-intuitive that an infinite and yet infinitely dense universe could be
created in a single instant at the Big Bang when R=0, but exactly that is predicted mathematically when k
does not equal 1. For comparison, an infinite plane has zero curvature but infinite area, whereas an infinite
cylinder is finite in one direction and a torus is finite in both. A toroidal universe could behave like a normal
universe with periodic boundary conditions, as seen in "wrap-around" video games such as Asteroids; a
traveler crossing an outer "boundary" of space going outwards would reappear instantly at another point on
the boundary moving inwards.
Prevailing model of the origin and expansion of spacetime and all that it contains.
The ultimate fate of the universe is still unknown, because it depends critically on the curvature index
k and the cosmological constant Λ. If the universe is sufficiently dense, k equals +1, meaning that its average
curvature throughout is positive and the universe will eventually recollapse in a Big Crunch, possibly starting
a new universe in a Big Bounce. Conversely, if the universe is insufficiently dense, k equals 0 or −1 and the
universe will expand forever, cooling off and eventually becoming inhospitable for all life, as the stars die and
all matter coalesces into black holes (the Big Freeze and the heat death of the universe). As noted above,
recent data suggests that the expansion speed of the universe is not decreasing as originally expected, but
increasing; if this continues indefinitely, the universe will eventually rip itself to shreds (the Big Rip).
Experimentally, the universe has an overall density that is very close to the critical value between recollapse
and eternal expansion; more careful astronomical observations are needed to resolve the question.

[edit] Big Bang model

Main articles: Big Bang, Timeline of the Big Bang, Nucleosynthesis, and Lambda-CDM model
The prevailing Big Bang model accounts for many of the experimental observations described above,
such as the correlation of distance and redshift of galaxies, the universal ratio of hydrogen:helium atoms, and
the ubiquitous, isotropic microwave radiation background. As noted above, the redshift arises from the metric
expansion of space; as the space itself expands, the wavelength of a photon traveling through space likewise
increases, decreasing its energy. The longer a photon has been traveling, the more expansion it has
undergone; hence, older photons from more distant galaxies are the most red-shifted. Determining the
correlation between distance and redshift is an important problem in experimental physical cosmology.
Chief nuclear reactions responsible for the relative abundances of light atomic nuclei observed
throughout the universe.
Other experimental observations can be explained by combining the overall expansion of space with
nuclear and atomic physics. As the universe expands, the energy density of the electromagnetic radiation
decreases more quickly than does that of matter, since the energy of a photon decreases with its wavelength.
Thus, although the energy density of the universe is now dominated by matter, it was once dominated by
radiation; poetically speaking, all was light. As the universe expanded, its energy density decreased and it
became cooler; as it did so, the elementary particles of matter could associate stably into ever larger
combinations. Thus, in the early part of the matter-dominated era, stable protons and neutrons formed, which
then associated into atomic nuclei. At this stage, the matter in the universe was mainly a hot, dense plasma
of negative electrons, neutral neutrinos and positive nuclei. Nuclear reactions among the nuclei led to the
present abundances of the lighter nuclei, particularly hydrogen, deuterium, and helium. Eventually, the
electrons and nuclei combined to form stable atoms, which are transparent to most wavelengths of radiation;
at this point, the radiation decoupled from the matter, forming the ubiquitous, isotropic background of
microwave radiation observed today.
Other observations are not answered definitively by known physics. According to the prevailing
theory, a slight imbalance of matter over antimatter was present in the universe's creation, or developed very
shortly thereafter, possibly due to the CP violation that has been observed by particle physicists. Although
the matter and antimatter mostly annihilated one another, producing photons, a small residue of matter
survived, giving the present matter-dominated universe. Several lines of evidence also suggest that a rapid
cosmic inflation of the universe occurred very early in its history (roughly 10−35 seconds after its creation).
Recent observations also suggest that the cosmological constant (Λ) is not zero and that the net mass-
energy content of the universe is dominated by a dark energy and dark matter that have not been
characterized scientifically. They differ in their gravitational effects. Dark matter gravitates as ordinary matter
does, and thus slows the expansion of the universe; by contrast, dark energy serves to accelerate the
universe's expansion.

[edit] Untestable proposals

[edit] Multiverse theory
Main articles: Multiverse, Many-worlds hypothesis, Bubble universe theory, and Parallel universe
Depiction of a multiverse of seven "bubble" universes, which are separate spacetime continua, each
having different physical laws, physical constants, and perhaps even different numbers of dimensions or
Some speculative theories have proposed that this universe is but one of a set of disconnected
universes, collectively denoted as the multiverse, altering the concept that the universe encompasses
everything.[15][73] By definition, there is no possible way for anything in one universe to affect another; if two
"universes" could affect one another, they would be part of a single universe. Thus, although some fictional
characters travel between parallel fictional "universes", this is, strictly speaking, an incorrect usage of the
term universe. The disconnected universes are conceived as being physical, in the sense that each should
have its own space and time, its own matter and energy, and its own physical laws — that also challenges the
definition of parallelity as these universes don't exist synchronously (since they have their own time) or in a
geometrically parallel way (since there's no interpretable relation between spatial positions of the different
universes). Such physically disconnected universes should be distinguished from the metaphysical
conception of alternate planes of consciousness, which are not thought to be physical places and are
connected through the flow of information. The concept of a multiverse of disconnected universes is very old;
for example, Bishop Étienne Tempier of Paris ruled in 1277 that God could create as many universes as he
saw fit, a question that was being hotly debated by the French theologians.[74]
There are two scientific senses in which multiple universes are discussed. First, disconnected
spacetime continua may exist; presumably, all forms of matter and energy are confined to one universe and
cannot "tunnel" between them. An example of such a theory is the chaotic inflation model of the early
universe.[75] Second, according to the many-worlds hypothesis, a parallel universe is born with every
quantum measurement; the universe "forks" into parallel copies, each one corresponding to a different
outcome of the quantum measurement. However, both senses of the term "multiverse" are speculative and
may be considered unscientific; no known experimental test in one universe could reveal the existence or
properties of another non-interacting universe.

[edit] Shape of the universe

For more details on this topic, see Shape of the universe.
The shape or geometry of the universe includes both local geometry in the observable universe and
global geometry, which we may or may not be able to measure. Shape can refer to curvature and topology.
More formally, the subject in practice investigates which 3-manifold corresponds to the spatial section in
comoving coordinates of the four-dimensional space-time of the universe. Analysis of data from WMAP
implies that the universe is spatially flat with only a 2% margin of error.[76]
Cosmologists normally work with a given space-like slice of spacetime called the comoving
coordinates. In terms of observation, the section of spacetime that can be observed is the backward light
cone (points within the cosmic light horizon, given time to reach a given observer). If the observable universe
is smaller than the entire universe (in some models it is many orders of magnitude smaller), one cannot
determine the global structure by observation: one is limited to a small patch.
In October 2001, NASA began collecting data with the Wilkinson Microwave Anisotropy Probe
(WMAP) on cosmic background radiation. Like visible light from distant stars and galaxies, cosmic
background radiation allows scientists to peer into the past to the time when the universe was in its infancy.
Density fluctuations in this radiation can also tell scientists much about the physical nature of space.[77]
NASA released the first WMAP cosmic background radiation data in February 2003. In 2009 the Planck
observatory was launched which will be able to analyze the microwave background at higher resolution,
providing more information on the shape of the early universe. The preliminary data will be released in
December 2010.

[edit] See also

Astronomy portal

Space portal

• Anthropic principle
• Big Bang
• Big Crunch
• Cosmic latte
• Cosmology
• Dyson's eternal intelligence
• Esoteric cosmology
• False vacuum
• Final anthropic principle
• Fine-tuned universe
• Heat death of the universe
• Hindu cycle of the universe
• Kardashev scale
• Multiverse
• Nucleocosmochronology
• Non-standard cosmology
• Omega Point
• Omniverse
• Rare Earth hypothesis
• Reality
• Shape of the universe
• Ultimate fate of the universe
• Vacuum genesis
• World view
• Zero-energy universe
[edit] Notes and references
1. ^ Webster's New World College Dictionary. Wiley Publishing, Inc.. 2010.
2. ^ The American Heritage® Dictionary of the English Language (4th ed.). Houghton Mifflin
Harcourt Publishing Company. 2010.
3. ^ Cambridge Advanced Learner's Dictionary.
4. ^ S. H. Suyu, P. J. Marshall, M. W. Auger, S. Hilbert, R. D. Blandford, L. V. E. Koopmans, C.
D. Fassnacht and T. Treu. Dissecting the Gravitational Lens B1608+656. II. Precision Measurements
of the Hubble Constant, Spatial Curvature, and the Dark Energy Equation of State. The Astrophysical
Journal, 2010; 711 (1): 201 DOI: 10.1088/0004-637X/711/1/201
5. ^ Lineweaver, Charles; Tamara M. Davis (2005). "Misconceptions about the Big Bang".
Scientific American.
03&page=5. Retrieved 2008-11-06.
6. ^ The Compact Edition of the Oxford English Dictionary , volume II, Oxford: Oxford University
Press, 1971, p.3518.
7. ^ a b Lewis and Short, A Latin Dictionary, Oxford University Press, ISBN 0-19-864201-6, pp.
1933, 1977–1978.
8. ^ Liddell and Scott, A Greek-English Lexicon, Oxford University Press, ISBN 0-19-864214-8,
9. ^ Liddell and Scott, pp.1345–1346.
10.^ Yonge, Charles Duke (1870). An English-Greek lexicon. New York: American Bok
Company. pp. 567.
11.^ Liddell and Scott, pp.985, 1964.
12.^ Lewis and Short, pp. 1881–1882, 1175, 1189–1190.
13.^ OED, pp. 909, 569, 3821–3822, 1900.
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18.^ Christian, Eric; Samar, Safi-Harb. "How large is the Milky Way?". Retrieved 2007-11-28.
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Galaxy". Astrophysical Journal 635: L37–L40. doi:10.1086/499161.
McConnachie, A. W.; Irwin, M. J.; Ferguson, A. M. N.; Ibata, R. A.; Lewis, G. F.; Tanvir, N. (2005).
"Distances and metallicities for 17 Local Group galaxies". Monthly Notices of the Royal Astronomical
Society 356 (4): 979–997. doi:10.1111/j.1365-2966.2004.08514.x.
20.^ Mackie, Glen (February 1, 2002). "To see the Universe in a Grain of Taranaki Sand".
Swinburne University. Retrieved 2006-12-20.
21.^ "Unveiling the Secret of a Virgo Dwarf Galaxy". ESO. 2000-05-03. Retrieved 2007-01-03.
22.^ "Hubble's Largest Galaxy Portrait Offers a New High-Definition View". NASA. 2006-02-28. Retrieved 2007-01-03.
23.^ "Star Count: ANU Astronomer makes best yet". 2003-07-17. Retrieved 2010-02-19.
24.^ N. Mandolesi, P. Calzolari, S. Cortiglioni, F. Delpino, G. Sironi (1986). "Large-scale
homogeneity of the Universe measured by the microwave background". Letters to Nature 319: 751–
753. doi:10.1038/319751a0.
25.^ Hinshaw, Gary (November 29, 2006). "New Three Year Results on the Oldest Light in the
Universe". NASA WMAP. Retrieved 2006-08-10.
26.^ Hinshaw, Gary (December 15, 2005). "Tests of the Big Bang: The CMB". NASA WMAP. Retrieved 2007-01-09.
27.^ Rindler (1977), p. 202.
28.^ Hinshaw, Gary (February 10, 2006). "What is the Universe Made Of?". NASA WMAP. Retrieved 2007-01-04.
29.^ "Five-Year Wilkinson Microwave Anisotropy Probe (WMAP) Observations: Data
Processing, Sky Maps, and Basic Results" (PDF).
Retrieved 2008-03-06.
30.^ Britt RR (2003-01-03). "Age of Universe Revised, Again". Retrieved 2007-01-08.
31.^ Wright EL (2005). "Age of the Universe". UCLA. Retrieved 2007-01-08.
Krauss LM, Chaboyer B (3 January 2003). "Age Estimates of Globular Clusters in the Milky Way:
Constraints on Cosmology". Science (American Association for the Advancement of Science) 299
(5603): 65–69. doi:10.1126/science.1075631. PMID 12511641.
ijkey=3D7y0Qonz=GO7ig.&keytype=3Dref&siteid=3Dsci. Retrieved 2007-01-08.
32.^ Wright, Edward L. (September 12, 2004). "Big Bang Nucleosynthesis". UCLA. Retrieved 2007-01-05.
M. Harwit, M. Spaans (2003). "Chemical Composition of the Early Universe". The Astrophysical
Journal 589 (1): 53–57. doi:10.1086/374415.
C. Kobulnicky, E. D. Skillman (1997). "Chemical Composition of the Early Universe". Bulletin of the
American Astronomical Society 29: 1329.
33.^ "Antimatter". Particle Physics and Astronomy Research Council. October 28, 2003. Retrieved 2006-08-10.
34.^ Landau and Lifshitz (1975), p.361.
35.^ WMAP Mission: Results – Age of the Universe
36.^ Luminet, Jean-Pierre; Boudewijn F. Roukema (1999). "Topology of the Universe: Theory
and Observations". Proceedings of Cosmology School held at Cargese, Corsica, August 1998 . Retrieved 2007-01-05.
Luminet, Jean-Pierre; J. Weeks, A. Riazuelo, R. Lehoucq, J.-P. Uzan (2003). "Dodecahedral space
topology as an explanation for weak wide-angle temperature correlations in the cosmic microwave
background" (subscription required). Nature 425 (6958): 593. doi:10.1038/nature01944.
PMID 14534579. Retrieved 2007-01-09.
37.^ Strobel, Nick (May 23, 2001). "The Composition of Stars". Astronomy Notes. Retrieved 2007-01-04.
"Have physical constants changed with time?". Astrophysics (Astronomy Frequently Asked
Questions). Retrieved 2007-01-04.
38.^ Stephen Hawking (1988). A Brief History of Time. Bantam Books. p. 125. ISBN 0-553-
39.^ Martin Rees (1999). Just Six Numbers. HarperCollins Publishers. ISBN 0-465-03672-4.
40.^ Adams, F.C. (2008). "Stars in other universes: stellar structure with different fundamental
constants". Journal of Cosmology and Astroparticle Physics 2008 (08): 010. doi:10.1088/1475-
41.^ Harnik, R.; Kribs, G.D. and Perez, G. (2006). "A universe without weak interactions".
Physical Review D 74: 035006. doi:10.1103/PhysRevD.74.035006.
42.^ Will Durant, Our Oriental Heritage:
"Two systems of Hindu thought propound physical theories suggestively similar to those
of Greece. Kanada, founder of the Vaisheshika philosophy, held that the world was
composed of atoms as many in kind as the various elements. The Jains more nearly
approximated to Democritus by teaching that all atoms were of the same kind,
producing different effects by diverse modes of combinations. Kanada believed light and
heat to be varieties of the same substance; Udayana taught that all heat comes from the
sun; and Vachaspati, like Newton, interpreted light as composed of minute particles
emitted by substances and striking the eye."
43.^ F. Th. Stcherbatsky (1930, 1962), Buddhist Logic, Volume 1, p.19, Dover, New York:
"The Buddhists denied the existence of substantial matter altogether. Movement
consists for them of moments, it is a staccato movement, momentary flashes of a
stream of energy... "Everything is evanescent“,... says the Buddhist, because there is no
stuff... Both systems [Sānkhya, and later Indian Buddhism] share in common a tendency
to push the analysis of Existence up to its minutest, last elements which are imagined as
absolute qualities, or things possessing only one unique quality. They are called
“qualities” (guna-dharma) in both systems in the sense of absolute qualities, a kind of
atomic, or intra-atomic, energies of which the empirical things are composed. Both
systems, therefore, agree in denying the objective reality of the categories of Substance
and Quality,... and of the relation of Inference uniting them. There is in Sānkhya
philosophy no separate existence of qualities. What we call quality is but a particular
manifestation of a subtle entity. To every new unit of quality corresponds a subtle
quantum of matter which is called guna “quality”, but represents a subtle substantive
entity. The same applies to early Buddhism where all qualities are substantive... or,
more precisely, dynamic entities, although they are also called dharmas ('qualities')."

44.^ a b c Craig, William Lane (June 1979). "Whitrow and Popper on the Impossibility of an
Infinite Past". The British Journal for the Philosophy of Science 30 (2): 165–170 [165–6].
45.^ Boyer, C. A History of Mathematics. Wiley, p. 54.
46.^ Otto E. Neugebauer (1945). "The History of Ancient Astronomy Problems and Methods",
Journal of Near Eastern Studies 4 (1), p. 1–38.
"the Chaldaean Seleucus from Seleucia"

47.^ George Sarton (1955). "Chaldaean Astronomy of the Last Three Centuries B. C.", Journal
of the American Oriental Society 75 (3), pp. 166–173 [169]:
"the heliocentrical astronomy invented by Aristarchos of Samos and still defended a
century later by Seleucos the Babylonian"

48.^ William P. D. Wightman (1951, 1953), The Growth of Scientific Ideas, Yale University Press
p.38, where Wightman calls him Seleukos the Chaldean.
49.^ Lucio Russo, Flussi e riflussi, Feltrinelli, Milano, 2003, ISBN 88-07-10349-4.
50.^ Bartel Leendert van der Waerden (1987), "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [527]
51.^ Bartel Leendert van der Waerden (1987), "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [527–9]
52.^ Bartel Leendert van der Waerden (1987). "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [529–34]
53.^ Bartel Leendert van der Waerden (1987). "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [534–7]
54.^ Nasr, Seyyed H. (1st edition in 1964, 2nd edition in 1993). An Introduction to Islamic
Cosmological Doctrines (2nd ed.). 1st edition by Harvard University Press, 2nd edition by State
University of New York Press. pp. 135–6. ISBN 0791415155.
55.^ Misner, Thorne and Wheeler (1973), p. 754.
56.^ Ragep, F. Jamil (2001a). "Tusi and Copernicus: The Earth's Motion in Context". Science in
Context (Cambridge University Press) 14 (1–2): 145–63.
57.^ Ragep, F. Jamil (2001b). "Freeing Astronomy from Philosophy: An Aspect of Islamic
Influence on Science". Osiris, 2nd Series 16 (Science in Theistic Contexts: Cognitive Dimensions):
49–64 & 66–71.
58.^ a b c d Misner, Thorne, and Wheeler (1973), p.755.
59.^ Misner, Thorne, and Wheeler (1973), p. 756.
60.^ de Cheseaux JPL (1744). Traité de la Comète. Lausanne. pp. 223ff. . Reprinted as
Appendix II in Dickson FP (1969). The Bowl of Night: The Physical Universe and Scientific Thought .
Cambridge, MA: M.I.T. Press. ISBN 978-0262540032.
61.^ Olbers HWM (1826). "Unknown title". Bode's Jahrbuch 111. . Reprinted as Appendix I in
Dickson FP (1969). The Bowl of Night: The Physical Universe and Scientific Thought . Cambridge,
MA: M.I.T. Press. ISBN 978-0262540032.
62.^ Jeans, J. H. (1902) Philosophical Transactions Royal Society of London, Series A , 199, 1.
63.^ Rindler, p. 196; Misner, Thorne, and Wheeler (1973), p. 757.
64.^ Misner, Thorne and Wheeler, p.756.
65.^ a b Einstein, A (1917). "Kosmologische Betrachtungen zur allgemeinen Relativitätstheorie".
Preussische Akademie der Wissenschaften, Sitzungsberichte 1917 (part 1): 142–152.
66.^ Rindler (1977), pp. 226–229.
67.^ Landau and Lifshitz (1975), pp. 358–359.
68.^ Einstein, A (1931). "Zum kosmologischen Problem der allgemeinen Relativitätstheorie".
Sitzungsberichte der Preussischen Akademie der Wissenschaften, Physikalisch-mathematische
Klasse 1931: 235–237.
Einstein A., de Sitter W. (1932). "On the relation between the expansion and the mean density of the
universe". Proceedings of the National Academy of Sciences 18 (3): 213–214.
doi:10.1073/pnas.18.3.213. PMID 16587663.
69.^ Hubble Telescope news release
70.^ BBC News story: Evidence that dark energy is the cosmological constant
71.^ Zel'dovich YB (1967). "Cosmological constant and elementary particles". Zh. Eksp. & Teor.
Fiz. Pis'ma 6: 883–884. English translation in Sov. Phys. — JTEP Lett., 6, pp. 316–317 (1967).
72.^ Friedmann A. (1922). "Über die Krümmung des Raumes". Zeitschrift für Physik 10: 377–
386. doi:10.1007/BF01332580.
73.^ Munitz MK (1959). "One Universe or Many?". Journal of the History of Ideas 12 (2): 231–
255. doi:10.2307/2707516.
74.^ Misner, Thorne and Wheeler (1973), p.753.
75.^ Linde A. (1986). "Eternal chaotic inflation". Mod. Phys. Lett. A1: 81.
Linde A. (1986). "Eternally existing self-reproducing chaotic inflationary universe". Phys. Lett. B175:
76.^ Shape of the Universe, WMAP website at NASA.

[edit] Further reading

• Landau, Lev, Lifshitz, E.M. (1975). The Classical Theory of Fields (Course of Theoretical
Physics, Vol. 2) (revised 4th English ed.). New York: Pergamon Press. pp. 358–397.
ISBN 9780080181769.
• Edward Robert Harrison (2000) Cosmology 2nd ed. Cambridge University Press. Gentle.
• Misner, C.W., Thorne, Kip, Wheeler, J.A. (1973). Gravitation. San Francisco: W. H. Freeman.
pp. 703–816. ISBN 978-0-7167-0344-0. The classic text for a generation.
• Rindler, W. (1977). Essential Relativity: Special, General, and Cosmological . New York:
Springer Verlag. pp. 193–244. ISBN 0-387-10090-3.
• Weinberg, S. (1993). The First Three Minutes: A Modern View of the Origin of the Universe
(2nd updated ed.). New York: Basic Books. ISBN 978-0465024377. OCLC 28746057. For lay
• -------- (2008) Cosmology. Oxford University Press. Challenging.
• Oscar Monchito (1987) Universe. What a concept. Colton, 23rd edition. For advanced
• Nussbaumer, Harry; Bieri, Lydia; Sandage, Allan (2009). Discovering the Expanding
Universe. Cambridge University Press. ISBN 978-0-521-51484-2.

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Pour les articles homonymes, voir Univers (homonymie).

Universum - C. Flammarion, gravure sur bois, Paris 1888, coloris : Heikenwaelder Hugo, Wien 1998.
L'Univers est l'ensemble de tout ce qui existe et les lois qui le régissent.
La cosmologie cherche à appréhender l'Univers d'un point de vue scientifique, comme l'ensemble de
la matière distribuée dans le temps et dans l'espace. Pour sa part, la cosmogonie vise à établir une théorie
de la création de l'Univers sur des bases philosophiques ou religieuses.
Selon le modèle standard, on ne connaît au plus que 5 % de la matière de l'Univers[1] ; le reste se
composerait de 25 % de matière noire et de 70 % d'énergie noire. Selon le modèle alternatif avec antimatière
soutenu notamment par Gabriel Chardin et Stephen Hawking, la quantité de matière présente dans l'univers
est 15 fois plus abondante que dans le modèle conventionnel[2].
• 1 Découverte de l'Univers dans l'Histoire
• 2 Naissance de l'Univers
• 2.1 L'expansion de l'Univers, son âge et le Big Bang
• 2.2 Taille de l'Univers, Univers observable
• 2.3 Forme de l'Univers
• 2.4 Modèle dimensionnel de l'Univers
• 3 Avenir de l'Univers
• 4 Notes et références
• 5 Voir aussi
• 5.1 Articles connexes
• 5.2 Liens externes

Découverte de l'Univers dans l'Histoire [modifier]

Articles détaillés : Monde (univers) et Révolution copernicienne.
Les Sciences grecques tentèrent de comprendre le monde et de l'expliquer :
• les philosophes Parménide, Platon, et Aristote avaient intégré l'idée d'une Terre sphérique,
mais ils la voyaient au centre de l'univers physique, alors que l'école de Milet se représentait la Terre
plate ;
• Ératosthène tenta de réaliser des calculs précis, notamment la mesure de la circonférence
d'un méridien terrestre ;
• Aristarque de Samos est le premier à envisager un modèle de système planétaire
héliocentré. Cette découverte ne fut alors pas suivie[3], pour des raisons philosophiques surtout
parce qu'une telle cosmologie est en désaccord avec la conception géocentrée du monde qui était
retenue par de grands philosophes comme Parménide, Platon, et Aristote. Il calcule aussi la distance
Terre-Lune pour laquelle il trouve une valeur discutée, mais qui se situe en tout état de cause dans
un ordre de grandeur acceptable[4], ainsi qu'une distance Terre-Soleil.[5] ;
• Hipparque poursuit ce travail : il recalcule, selon des méthodes nouvelles, la distance Terre-
Soleil ainsi que distance Terre-Lune (pour la quelle il retient la valeur de 67 1/3 rayons terrestres,
contre 60,2 en réalité[6]), recense 1 500 étoiles[réf. nécessaire], retrouve approximativement la
période de précession des équinoxes, qui était déjà connue des Babyloniens.[réf. nécessaire]
• Ptolémée poursuit le travail d'Hipparque. Son Almageste sera la référence astronomique
essentielle pendant treize siècles.
L'Univers selon le système de Ptolémée, vu par Andreas Cellarius en 1660/61.
Ces connaissances du monde grec perdureront et influenceront les sciences arabes après
l'effondrement de l'Empire romain d'Occident. Elles resteront présentes en Orient (particulièrement, avec des
hauts et des bas, à Byzance[7]), même si Cosmas d'Alexandrie tente, sans succès, de restaurer le modèle
d'un monde plat.
La Renaissance porte à son apogée cette représentation du monde, grâce aux explorations et aux
grandes découvertes qui eurent lieu du XIIIe au XVIe siècles, à partir de systèmes géographiques et
cosmologiques très élaborés (projection de Mercator).
La révolution copernicienne bouleverse cette cosmologie en trois étapes :
1. Copernic redécouvre l'héliocentrisme. Toutefois, cette redécouverte n'est que partiellement
révolutionnaire : en effet, Copernic reste attaché aux sphères transparentes du modèle d'Aristote
(pourtant délaissé par Ptolémée) censées soutenir les planètes et leur imprimer leur mouvement ; il
présente son système comme un simple artifice destiné à simplifier les calculs, ce qui lui évite des
ennuis avec le clergé.
2. Le dominicain Giordano Bruno défend la réalité du modèle héliocentrique et l'étend à toutes
les étoiles, ouvrant la dimension de l'univers physique à l'infini. Pour cette raison, entre autres, il sera
brulé au bûcher en tant qu'hérétique.
3. Kepler, Galilée et Newton posent les bases fondamentales de la mécanique à partir du
mouvement des planètes, grâce à leurs études respectivement du mouvement elliptique des
planètes autour du Soleil, l'affinement des observations astronomiques avec la définition du
mouvement uniformément accéléré, et la formalisation mathématique de la force de gravité.
L'Univers, toutefois, reste confiné dans le système solaire.
Des modèles physiques tels que la sphère armillaire ou l'astrolabe ont été élaborés. Ils permettent
d'enseigner et de calculer la position des astres dans le ciel visible. Aujourd'hui encore, la carte du ciel
mobile aide les astronomes amateurs à se repérer dans le ciel, c'est une ré-incarnation de l'astrolabe.

Naissance de l'Univers [modifier]

L'expansion de l'Univers, son âge et le Big Bang [modifier]

Articles détaillés : Frise chronologique du Big Bang, Expansion de l'Univers et Big Bang.
Les observations du décalage vers le rouge des rayonnements électromagnétiques en provenance
d'autres galaxies suggèrent que celles-ci s'éloignent de notre galaxie, à une vitesse radiale d'éloignement
supposée proportionnelle à ce décalage.
En étudiant les galaxies proches, Edwin Hubble s'est aperçu que la vitesse d'éloignement d'une
galaxie était proportionnelle à sa distance par rapport à l'observateur (loi de Hubble); une telle loi correspond
à un Univers visible en expansion.
Bien que la constante de Hubble ait été révisée par le passé dans d'importantes proportions (dans
un rapport de 10 à 1), la loi de Hubble a été extrapolée aux galaxies éloignées, pour lesquelles la distance
ne peut être calculée au moyen de la parallaxe ; cette loi est ainsi utilisée pour déterminer la distance des
galaxies les plus lointaines.
En extrapolant l'expansion de l'Univers dans le passé, on arrive à une époque où celui-ci a dû être
beaucoup plus chaud et beaucoup plus dense qu'aujourd'hui. C'est le modèle du Big Bang qui est un
ingrédient essentiel du modèle standard de la cosmologie actuelle et possède aujourd'hui un grand nombre
de confirmations expérimentales. La description du début de l'histoire de l'Univers par ce modèle ne
commence cependant qu'après qu'il fût sorti d'une période appelée ère de Planck durant laquelle l'échelle
d'énergie de l'Univers était si grande que le modèle standard n'est pas en mesure de décrire les
phénomènes quantiques qui s'y sont déroulés. Durant cette époque, seule une théorie de la gravitation
quantique pourrait expliquer le comportement microscopique de la matière sous l'influence importante de la
gravité. Mais les physiciens ne disposent pas encore (en 2010) d'une telle théorie. Pour des raisons de
cohérence avec les observations, après l'ère de Planck le modèle du Big Bang privilégie aujourd'hui
l'existence d'une phase d'inflation cosmique très brève mais durant laquelle l'Univers aurait grandi de façon
extrêmement rapide. C'est suite à cette phase que l'essentiel des particules de l'Univers auraient été créées
avec une haute température, enclenchant un grand nombre de processus importants[8] qui ont finalement
abouti à l'émission d'une grande quantité de lumière, appelé fond diffus cosmologique, qui peut être
aujourd'hui observé avec une grande précision par toute une série d'instruments (ballons-sondes, sondes
C'est l'observation de ce rayonnement fossile micro-onde, remarquablement uniforme dans toutes
les directions qui constitue aujourd'hui l'élément capital qui assoit le modèle du Big Bang comme description
correcte de l'Univers dans son passé lointain. Beaucoup d'éléments du modèle devraient encore être
affinés[9], mais il y a aujourd'hui consensus de la communauté scientifique autour du modèle du Big Bang.
Dans le cadre du modèle ΛCDM, qui est le plus simple incorporant tous les éléments que l'on vient
d'évoquer, les contraintes issues des observations de la sonde WMAP[10] sur les paramètres
cosmologiques indiquent une valeur la plus probable pour l'âge de l'Univers à environ 13,7 milliards
d'années[11] avec une incertitude de 0,2 milliard d'années, ce qui est en accord avec les données
indépendantes issues de l'observation des amas globulaires[12] ainsi que celle des naines blanches[13].

Taille de l'Univers, Univers observable [modifier]

Article détaillé : Univers observable.
À ce jour, rien ne nous permet de confirmer que l'Univers est soit fini, soit infini. Certains théoriciens
penchent pour un Univers infini, d'autres pour un Univers fini mais non borné.
Les articles populaires et professionnels de recherche en cosmologie emploient souvent le terme
« Univers » dans le sens d'« Univers observable ». Nous vivons au centre de l'Univers observable, en
contradiction apparente avec le principe de Copernic qui dit que l'Univers est plus ou moins uniforme et ne
possède aucun centre en particulier. C'est simplement parce que la lumière ne se déplace pas à une vitesse
infinie et que les observations que nous faisons proviennent donc du passé. En effet, en regardant de plus
en plus loin, nous voyons des choses qui se sont passées à une époque de plus en plus proche du Big-
Bang. Et puisque la lumière se déplace à la même vitesse dans toutes les directions, tous les observateurs
vivent au centre de leur Univers observable (sur Terre, nous avons pratiquement tous le même). On appelle
« horizon cosmologique » la première lumière émise par le Big-Bang il y a 13,7 milliards d'années. Il nous est
impossible de voir plus loin.
On estime que le diamètre de cet Univers observable est de 100 milliards d'années lumière[14].
Celui-ci contient environ 7×1022 étoiles, répandues dans environ 1011 galaxies, elles-mêmes organisées en
amas et superamas de galaxies[14]. Mais le nombre de galaxies pourrait être encore plus grand, selon le
champ profond observé avec le télescope spatial Hubble.
Il est cependant probable que l'Univers que nous voyons n'est qu'une infime partie d'un Univers réel
beaucoup plus grand. Selon les derniers modèles cosmologiques, la taille minimale de l'Univers réel serait

de [Quoi ?] soit un 1 suivi de dix milliards de zéros[14]. (A priori, l'unité n'a pas d'importance:
métre, millimetre, kilometre ou UA. Ca représente seulement quelques zéro en plus ou en moins...)
On pourrait raisonner que, l'Univers contenant par définition tout ce qui existe, y compris l'espace-
temps (et c'est une précision essentielle), il ne peut avoir de « bord » tel que nous concevons intuitivement
cette notion. En effet, l'existence d'un bord impliquerait qu'au-delà de ce bord, on ne serait plus dans
l'Univers, ce qui serait absurde. Mais si l'Univers n'a pas de bord au sens intuitif de ce terme, alors son
expansion n'est pas intuitive non plus : si elle l'était, dans quoi l'Univers serait-il en expansion ?
On voit ici les apparents paradoxes entraînés par l'utilisation de notions dites « intuitives »... qui ne
sont que le reflet d'une perception locale de la réalité. Donner une définition précise à ces notions intuitives
permet de faire disparaître ces paradoxes. On voit que ce problème échappe à nos raisonnements
simplistes, qui se fondent sur des hypothèses fausses telles que « l'Univers est galiléen », ou « un espace
courbe est nécessairement inclus dans un espace de dimension supérieure ». En définitive, une définition
précise de la notion d'Univers permet de tenter de résoudre ces apparents paradoxes. Les paradoxes
restants permettent de démontrer... que la définition donnée à l'Univers n'a pas sens. On sait en fait
aujourd'hui qu'une définition, même formelle, d'un ensemble n'implique pas son existence.
En bref, le mot « Univers » reste à définir.

Forme de l'Univers [modifier]

Articles détaillés : Topologie de l'Univers et Courbure spatiale.
Une importante question de cosmologie qui reste sans réponse est la topologie de l'Univers.
1. Est-ce que l'Univers est « plat » ? C'est-à-dire : est-ce que le théorème de Pythagore pour
les triangles droits est valide à de plus grandes échelles ? Actuellement, la plupart des cosmologues
croient que l'Univers observable est (presque) plat, juste comme la Terre est (presque) plate.
2. Est-ce que l'Univers est simplement connexe ? Selon le modèle standard du Big Bang,
l'Univers n'a aucune frontière spatiale, mais peut néanmoins être de taille finie.
Ceci peut être compris par une analogie bidimensionnelle : la surface de la Terre n'a aucun bord,
mais possède une aire bien déterminée.
Vous pouvez également penser à un cylindre et imaginer de coller les 2 extrémités du cylindre ensemble,
mais sans plier le cylindre.
C'est aussi un espace bidimensionnel avec une surface finie, mais au contraire de la surface de la Terre, il
est plat, et peut ainsi servir de meilleur modèle.
Par conséquent, à proprement parler, nous devrions appeler les étoiles et les galaxies mentionnées
ci-dessus « images » d'étoiles et de galaxies, puisqu'il est possible que l'Univers soit fini et si petit que nous
pouvons voir une ou plusieurs fois autour de lui, et le vrai nombre d'étoiles et de galaxies physiquement
distinctes pourrait être plus petit. Des hypothèses d'Univers multiconnexe ont été proposées et sont en cours

Modèle dimensionnel de l'Univers [modifier]

L’Univers a-t-il 3, 6, 10 dimensions ou plus ?
La théorie des cordes prédit qu’espace et matière sont consubstantiels. Il n’y a pas de « contenant »
(l’espace) mais un fond d’espace-temps qui interagit avec la matière. Dans certains cas particuliers, la notion
de « nombre de dimensions de l’espace » dépend de l’intensité avec laquelle les cordes réagissent entre
elles. Si cette interaction est faible, elles semblent se propager dans un espace à neuf dimensions -
auxquelles il faut rajouter celle du temps. Si cette interaction croît, cela développe une dimension de plus (ou
plus en fonction de l’intensité de l’interaction) à laquelle il faut toujours rajouter celle du temps. Supposons
maintenant qu’on enferme l’Univers dans un espace « fini » (une boîte pour être concret) et que cet espace
rapetisse jusqu’à 10-32 centimètre de côté, la théorie des cordes le prédit équivalent à un Univers très grand.
La conception d’espace est fondamentalement bouleversée. La mise en route du grand collisionneur de
hadrons de Genève, Large Hadron Collider (plus communément appelé LHC), viendra peut être confirmer
cette théorie. Elle ne pourra en revanche pas l'infirmer, car aucun ordre de grandeur n'a été prédit par la
théorie des cordes. Ainsi, si le phénomène n'est pas détecté, cela pourrait signifier que trop peu d'énergie a
été générée pour rendre le phénomène observable, sans impliquer pour autant que la théorie soit
nécessairement erronée.
Avenir de l'Univers [modifier]
Les objets galactiques auront une fin : le Soleil, par exemple, s'éteindra dans 5 (à 7) milliards
d'années, lorsqu'il aura consumé tout son combustible. À terme, les autres étoiles suivront elles aussi dans
des cataclysmes cosmologiques (explosions, effondrements). Déjà les naissances d'étoiles se
ralentissent[15] faute de matière, qui se raréfie au fil du temps. Dans 20 milliards d'années environ, aucun
astre ne s'allumera plus. L'Univers sera peuplé d'étoiles éteintes (étoiles à neutrons, naines blanches, trous
noirs) et des naines rouges résiduelles. À bien plus longues échéances, les galaxies se désagrègeront dans
des collisions géantes par leurs interactions gravitationnelles internes et externes[16].
En ce qui concerne le contenant (l'espace), certains[Qui ?] pensent que le processus d'expansion
sera gravitationnellement ralenti et s'inversera selon le scénario du Big Crunch[17]. Pour d'autres[Qui ?],
l'expansion, qui semble à présent accélérée par la présence d'une énergie répulsive de nature inconnue
(l'énergie sombre), continuera à jamais. Peu à peu, les astres éteints s'agglutineront en trous noirs.
L'Univers, sans aucune structure, ne sera plus qu'un bain de photons de plus en plus froids[18]. Toute
activité dans l'Univers s'éteindra ainsi à jamais. Si au contraire la quantité d'énergie sombre croît, l'Univers
continuera son expansion à une vitesse toujours plus grande pour exploser à toutes les échelles : toute la
matière qui le compose (y compris les atomes) se déchirera par dilatation de l'espace. C'est le Big Rip
(littéralement : « grand déchirement »). Certains modèles prévoient une telle fin dans 22 milliards d'années.
Chacun de ces scénarios dépend donc de la quantité d'énergie sombre que contiendra l'Univers à un
moment donné. Actuellement, l'état des connaissances suggère non seulement qu'il y a insuffisamment de
masse et d'énergie pour provoquer ce Big Rip, mais que l'expansion de l'Univers semble s'accélérer et
continuera donc pour toujours[réf. nécessaire].
Notes et références [modifier]
1. ↑ NASA WMAP What is the universe made of ? [archive]
2. ↑ Antimatière : est-elle la clé de l'Univers ?, Science et vie, octobre 2009, n°1105, p.60.
3. ↑ À notre connaissance, un seul autre savant de l'antiquité fut de cet avis, Séleucos de
4. ↑ Otto Neugebauer, A history of ancient mathematical astronomy, Berlin ; New York :
Springer-Verlag, 1975, p. 634 ss. Aristarque ne donne pas le résultat de ses calculs, mais de ses
données (diamètre apparent angulaire de la Lune : 2° ; diamètre de la Lune : 1/3 de diamètre
lunaire), on peut déduire une distance Terre-Lune de 40 rayons terrestres environ, contre 60,2 en
réalité. Mais Neugebauer estime que c'est un angle de 1/2° et non de 2° qu'Aristarque tenait pour
correct, ce qui aboutirait à 80 rayons terrestres pour la distance Terre-Lune. Voir Aristarque.
5. ↑ Neugebauer, Loc. cit..
6. ↑ Cf. Des grandeurs et des distances du Soleil et de la Lune.
7. ↑ Cf. sciences grecques.
8. ↑ Comme la nucléosynthèse primordiale par exemple ou encore la baryogénèse.
9. ↑ Il n'est pas encore possible de bien discriminer entre plusieurs modèles d'inflation pour
donner un exemple.
10.↑ Lancée par la NASA.
11.↑ (en) D.N. Spergel et al., Wilkinson microwave anisotropy probe (wmap) three year
results : implications for cosmology. [archive] soumis à Astrophys. J., prépublication disponible sur la
base de données arXiv.
12.↑ (en) Chaboyer, B. & Krauss, Theoretical Uncertainties in the Subgiant--Mass Age
Relation and the Absolute Age of Omega Cen [archive]L. M. 2002, ApJ, 567, L45.
13.↑ (en)B. Hanser et al HST Observations of the White Dwarf Cooling Sequence of
M4 [archive] The Astrophysical Journal Supplement Series, Volume 155, Issue 2, p. 551-576.,
prépublication disponible sur l'arXiv.
14.↑ a, b et c Science-et-Vie Hors-Série n°242, mars 2008. "L'Univers en chiffres".
15.↑ Actuellement, l'observation de notre galaxie dénombre la naissance d'une ou deux étoiles
par an.
16.↑ Jean-Pierre Luminet, Astrophysicien, CNRS, Observatoire de Paris-Meudon, in Sciences
& Avenir N°729, Novembre 2007.
17.↑ Littéralement : « grand écrasement ».
18.↑ D'après une théorie de Stephen Hawking (dans son livre Une brève histoire du temps), si
l'Univers continue indéfiniment à s'étendre, les particules issues d'explosions successives ne seront
plus assez proches les unes des autres pour recréer des étoiles après leur explosion.

Voir aussi [modifier]

Sur les autres projets Wikimédia :
• Univers sur le Wiktionnaire (dictionnaire universel)
• Univers sur Wikibooks (livres pédagogiques)
• Univers sur Wikisource (bibliothèque universelle)
• Modèle cosmologique
• Cosmologie non standard
Articles connexes [modifier]
• Voie lactée
• Monde (univers)
• Radiotélescope
• Observatoire astronomique
• Destin de l'Univers
• Âge de l'Univers
• Masse de l'Univers
• Théorie d'Everett
• Table des constantes astrophysiques
• Densité critique
• Énergie sombre
• Matière noire

Liens externes [modifier]

• (fr) Univers : page de l'Institut national des sciences de l'Univers

• Portail de l’astronomie

• Portail de la cosmologie

From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is about the planet. For other uses, see Earth (disambiguation).

"Blue Marble" photograph of Earth,
taken from Apollo 17
Pronunciation i /ˈɜrθ/

Adjective earthly, tellurian, telluric, terran, terrestrial.

Orbital characteristics
Epoch J2000.0[note 1]

152,098,232 km
1.01671388 AU[note 2]

147,098,290 km
0.98329134 AU[note 2]

149,598,261 km
Semi-major axis
1.00000261 AU[1]

Eccentricity 0.01671123[1]

365.256363004 days[2]
Orbital period
1.000017421 yr

29.78 km/s[3]
Average orbital speed
107,200 km/h

7.155° to Sun's equator

1.57869°[4] to invariable plane

Longitude of ascending node 348.73936°[3][note 3]

Argument of perihelion 114.20783°[3][note 4]

Satellites 1 (the Moon)

Physical characteristics
Mean radius 6,371.0 km[5]

Equatorial radius 6,378.1 km[6]

Polar radius 6,356.8 km[7]

Flattening 0.0033528[6]

40,075.16 km (equatorial)[8]
40,008.00 km (meridional)[8]

510,072,000 km2[9][10][note 5]
Surface area 148,940,000 km2 land (29.2 %)
361,132,000 km2 water (70.8 %)

Volume 1.08321 × 1012 km3[3]

Mass 5.9736 × 1024 kg[3]

Mean density 5.515 g/cm3[3]

Equatorial surface gravity 9.780327 m/s2[11]

0.99732 g

Escape velocity 11.186 km/s[3]

Sidereal rotation 0.99726968 d[12]

period 23h 56m 4.100s

Equatorial rotation velocity 1,674.4 km/h (465.1 m/s)[13]

Axial tilt 23°26'21".4119[2]

Albedo 0.367[3]

Surface temp.
min mean max
184 K[14 287.2 K[15 331 K[16
] ] ]
-89.2 °C 14 °C 57.8 °C
Surface pressure 101.325 kPa (MSL)

78.08% nitrogen (N2)[3]

20.95% oxygen (O2)
Composition 0.93% argon
0.038% carbon dioxide
About 1% water vapor (varies with climate)
Earth (or the Earth) is the third planet from the Sun, and the densest and fifth-largest of the eight
planets in the Solar System. It is also the largest of the Solar System's four terrestrial planets. It is sometimes
referred to as the World, the Blue Planet,[note 6] or by its Latin name, Terra.[note 7]
Home to millions of species[17] including humans, Earth is currently the only place in the universe
where life is known to exist. The planet formed 4.54 billion years ago,[18] and life appeared on its surface
within a billion years. Earth's biosphere has significantly altered the atmosphere and other abiotic conditions
on the planet, enabling the proliferation of aerobic organisms as well as the formation of the ozone layer
which, together with Earth's magnetic field, blocks harmful solar radiation, permitting life on land.[19] The
physical properties of the Earth, as well as its geological history and orbit, have allowed life to persist during
this period. The planet is expected to continue supporting life for at least another 500 million years.[20][21]
Earth's outer surface is divided into several rigid segments, or tectonic plates, that migrate across the
surface over periods of many millions of years. About 71% of the surface is covered with salt water oceans,
the remainder consisting of continents and islands which together have many lakes and other sources of
water contributing to the hydrosphere. Liquid water, necessary for all known life, is not known to exist on any
other planet's surface.[note 8][note 9] Earth's poles are mostly covered with solid ice (Antarctic ice sheet) or
sea ice (Arctic ice cap). The planet's interior remains active, with a thick layer of relatively solid mantle, a
liquid outer core that generates a magnetic field, and a solid iron inner core.
Earth interacts with other objects in space, especially the Sun and the Moon. At present, Earth orbits
the Sun once for every roughly 366.26 times it rotates about its axis, which is equal to 365.26 solar days, or
one sidereal year.[note 10] The Earth's axis of rotation is tilted 23.4° away from the perpendicular to its orbital
plane,[22] producing seasonal variations on the planet's surface with a period of one tropical year (365.24
solar days). Earth's only known natural satellite, the Moon, which began orbiting it about 4.53 billion years
ago, provides ocean tides, stabilizes the axial tilt and gradually slows the planet's rotation. Between
approximately 3.8 billion and 4.1 billion years ago, numerous asteroid impacts during the Late Heavy
Bombardment caused significant changes to the greater surface environment.
Both the mineral resources of the planet, as well as the products of the biosphere, contribute
resources that are used to support a global human population. These inhabitants are grouped into about 200
independent sovereign states, which interact through diplomacy, travel, trade, and military action. Human
cultures have developed many views of the planet, including personification as a deity, a belief in a flat Earth
or in Earth as the center of the universe, and a modern perspective of the world as an integrated environment
that requires stewardship.
• 1 Chronology
• 1.1 Evolution of life
• 1.2 Future
• 2 Composition and structure
• 2.1 Shape
• 2.2 Chemical composition
• 2.3 Internal structure
• 2.4 Heat
• 2.5 Tectonic plates
• 2.6 Surface
• 2.7 Hydrosphere
• 2.8 Atmosphere
• 2.8.1 Weather and climate
• 2.8.2 Upper atmosphere
• 2.9 Magnetic field
• 3 Orbit and rotation
• 3.1 Rotation
• 3.2 Orbit
• 3.3 Axial tilt and seasons
[edit] Chronology
Main article: History of the Earth
See also: Geological history of Earth
Scientists have been able to reconstruct detailed information about the planet's past. The earliest
dated Solar System material is dated to 4.5672 ± 0.0006 billion years ago,[23] and by 4.54 billion years ago
(within an uncertainty of 1%)[18] the Earth and the other planets in the Solar System had formed out of the
solar nebula—a disk-shaped mass of dust and gas left over from the formation of the Sun. This assembly of
the Earth through accretion was thus largely completed within 10–20 million years.[24] Initially molten, the
outer layer of the planet Earth cooled to form a solid crust when water began accumulating in the
atmosphere. The Moon formed shortly thereafter, 4.53 billion years ago.[25]
The current consensus model[26] for the formation of the Moon is the giant impact hypothesis, in
which the Moon formed as a result of a Mars-sized object (sometimes called Theia) with about 10% of the
Earth's mass[27] impacting the Earth in a glancing blow.[28] In this model, some of this object's mass would
have merged with the Earth and a portion would have been ejected into space, but enough material would
have been sent into orbit to form the Moon.
Outgassing and volcanic activity produced the primordial atmosphere. Condensing water vapor,
augmented by ice and liquid water delivered by asteroids and the larger proto-planets, comets, and trans-
Neptunian objects produced the oceans.[29] The newly formed Sun was only 70% of its present luminosity,
yet evidence shows that the early oceans remained liquid—a contradiction dubbed the faint young Sun
paradox. A combination of greenhouse gases and higher levels of solar activity served to raise the Earth's
surface temperature, preventing the oceans from freezing over.[30] By 3.5 billion years ago, the Earth's
magnetic field was established, which helped prevent the atmosphere from being stripped away by the solar
Two major models have been proposed for the rate of continental growth:[32] steady growth to the
present-day[33] and rapid growth early in Earth history.[34] Current research shows that the second option is
most likely, with rapid initial growth of continental crust[35] followed by a long-term steady continental area.
[36][37][38] On time scales lasting hundreds of millions of years, the surface continually reshaped as
continents formed and broke up. The continents migrated across the surface, occasionally combining to form
a supercontinent. Roughly 750 million years ago (Ma), one of the earliest known supercontinents, Rodinia,
began to break apart. The continents later recombined to form Pannotia, 600–540 Ma, then finally Pangaea,
which broke apart 180 Ma.[39]

[edit] Evolution of life

Main article: Evolutionary history of life
At present, Earth provides the only example of an environment that has given rise to the evolution of
life.[40] Highly energetic chemistry is believed to have produced a self-replicating molecule around
4 billion years ago and half a billion years later the last common ancestor of all life existed.[41] The
development of photosynthesis allowed the Sun's energy to be harvested directly by life forms; the resultant
oxygen accumulated in the atmosphere and formed a layer of ozone (a form of molecular oxygen [O3]) in the
upper atmosphere. The incorporation of smaller cells within larger ones resulted in the development of
complex cells called eukaryotes.[42] True multicellular organisms formed as cells within colonies became
increasingly specialized. Aided by the absorption of harmful ultraviolet radiation by the ozone layer, life
colonized the surface of Earth.[43]
Since the 1960s, it has been hypothesized that severe glacial action between 750 and 580 Ma,
during the Neoproterozoic, covered much of the planet in a sheet of ice. This hypothesis has been termed
"Snowball Earth", and is of particular interest because it preceded the Cambrian explosion, when
multicellular life forms began to proliferate.[44]
Following the Cambrian explosion, about 535 Ma, there have been five major mass extinctions.[45]
The most recent such event was 65 Ma, when an asteroid impact triggered the extinction of the (non-avian)
dinosaurs and other large reptiles, but spared some small animals such as mammals, which then resembled
shrews. Over the past 65 million years, mammalian life has diversified, and several million years ago, an
African ape-like animal such as Orrorin tugenensis gained the ability to stand upright.[46] This enabled tool
use and encouraged communication that provided the nutrition and stimulation needed for a larger brain. The
development of agriculture, and then civilization, allowed humans to influence the Earth in a short time span
as no other life form had,[47] affecting both the nature and quantity of other life forms.
The present pattern of ice ages began about 40 Ma and then intensified during the Pleistocene about
3 Ma. High-latitude regions have since undergone repeated cycles of glaciation and thaw, repeating every
40–100,000 years. The last continental glaciation ended 10,000 years ago.[48]

[edit] Future
Main article: Future of the Earth
See also: Risks to civilization, humans and planet Earth
The life cycle of the Sun
The future of the planet is closely tied to that of the Sun. As a result of the steady accumulation of
helium at the Sun's core, the star's total luminosity will slowly increase. The luminosity of the Sun will grow by
10% over the next 1.1 Gyr (1.1 billion years) and by 40% over the next 3.5 Gyr.[49] Climate models indicate
that the rise in radiation reaching the Earth is likely to have dire consequences, including the loss of the
planet's oceans.[50]
The Earth's increasing surface temperature will accelerate the inorganic CO2 cycle, reducing its
concentration to levels lethally low for plants (10 ppm for C4 photosynthesis) in approximately 500 million[20]
to 900 million years. The lack of vegetation will result in the loss of oxygen in the atmosphere, so animal life
will become extinct within several million more years.[51] After another billion years all surface water will
have disappeared[21] and the mean global temperature will reach 70 °C[51] (158 °F). The Earth is expected
to be effectively habitable for about another 500 million years from that point,[20] although this may be
extended up to 2.3 billion years if the nitrogen is removed from the atmosphere.[52] Even if the Sun were
eternal and stable, the continued internal cooling of the Earth would result in a loss of much of its CO 2 due to
reduced volcanism,[53] and 35% of the water in the oceans would descend to the mantle due to reduced
steam venting from mid-ocean ridges.[54]
The Sun, as part of its evolution, will become a red giant in about 5 Gyr. Models predict that the Sun
will expand out to about 250 times its present radius, roughly 1 AU (150,000,000 km).[49][55] Earth's fate is
less clear. As a red giant, the Sun will lose roughly 30% of its mass, so, without tidal effects, the Earth will
move to an orbit 1.7 AU (250,000,000 km) from the Sun when the star reaches it maximum radius. The
planet was therefore initially expected to escape envelopment by the expanded Sun's sparse outer
atmosphere, though most, if not all, remaining life would have been destroyed by the Sun's increased
luminosity (peaking at about 5000 times its present level).[49] However, a more recent simulation indicates
that Earth's orbit will decay due to tidal effects and drag, causing it to enter the red giant Sun's atmosphere
and be vaporized.[55]

[edit] Composition and structure

Main article: Earth science
Further information: Earth physical characteristics tables
Earth is a terrestrial planet, meaning that it is a rocky body, rather than a gas giant like Jupiter. It is
the largest of the four solar terrestrial planets in size and mass. Of these four planets, Earth also has the
highest density, the highest surface gravity, the strongest magnetic field, and fastest rotation.[56] It also is the
only terrestrial planet with active plate tectonics.[57]

[edit] Shape
Main article: Figure of the Earth

Size comparison of inner planets (left to right): Mercury, Venus, Earth and Mars
The shape of the Earth is very close to that of an oblate spheroid, a sphere flattened along the axis
from pole to pole such that there is a bulge around the equator.[58] This bulge results from the rotation of the
Earth, and causes the diameter at the equator to be 43 km larger than the pole to pole diameter.[59] The
average diameter of the reference spheroid is about 12,742 km, which is approximately 40,000 km/π, as the
meter was originally defined as 1/10,000,000 of the distance from the equator to the North Pole through
Paris, France.[60]
Local topography deviates from this idealized spheroid, though on a global scale, these deviations
are very small: Earth has a tolerance of about one part in about 584, or 0.17%, from the reference spheroid,
which is less than the 0.22% tolerance allowed in billiard balls.[61] The largest local deviations in the rocky
surface of the Earth are Mount Everest (8848 m above local sea level) and the Mariana Trench (10,911 m
below local sea level). Because of the equatorial bulge, the surface locations farthest from the center of the
Earth are the summits of Mount Chimborazo in Ecuador and Huascarán in Peru.[62][63][64]
Chemical composition of the crust[65]
Compound Formula
Continental Oceanic

silica SiO2 60.2% 48.6%

alumina Al2O3 15.2% 16.5%

lime CaO 5.5% 12.3%

magnesia MgO 3.1% 6.8%

iron(II) oxide FeO 3.8% 6.2%

sodium oxide Na2O 3.0% 2.6%

potassium oxide K2O 2.8% 0.4%

iron(III) oxide Fe2O3 2.5% 2.3%

water H2O 1.4% 1.1%

carbon dioxide CO2 1.2% 1.4%

titanium dioxide TiO2 0.7% 1.4%

phosphorus pentoxide P2O5 0.2% 0.3%

Total 99.6% 99.9%

[edit] Chemical composition

See also: Abundance of elements on Earth
The mass of the Earth is approximately 5.98 × 1024 kg. It is composed mostly of iron (32.1%), oxygen
(30.1%), silicon (15.1%), magnesium (13.9%), sulfur (2.9%), nickel (1.8%), calcium (1.5%), and aluminium
(1.4%); with the remaining 1.2% consisting of trace amounts of other elements. Due to mass segregation, the
core region is believed to be primarily composed of iron (88.8%), with smaller amounts of nickel (5.8%),
sulfur (4.5%), and less than 1% trace elements.[66]
The geochemist F. W. Clarke calculated that a little more than 47% of the Earth's crust consists of
oxygen. The more common rock constituents of the Earth's crust are nearly all oxides; chlorine, sulfur and
fluorine are the only important exceptions to this and their total amount in any rock is usually much less than
1%. The principal oxides are silica, alumina, iron oxides, lime, magnesia, potash and soda. The silica
functions principally as an acid, forming silicates, and all the commonest minerals of igneous rocks are of this
nature. From a computation based on 1,672 analyses of all kinds of rocks, Clarke deduced that 99.22% were
composed of 11 oxides (see the table at right). All the other constituents occur only in very small quantities.

[edit] Internal structure

Main article: Structure of the Earth
The interior of the Earth, like that of the other terrestrial planets, is divided into layers by their
chemical or physical (rheological) properties. The outer layer of the Earth is a chemically distinct silicate solid
crust, which is underlain by a highly viscous solid mantle. The crust is separated from the mantle by the
Mohorovičić discontinuity, and the thickness of the crust varies: averaging 6 km under the oceans and 30–
50 km on the continents. The crust and the cold, rigid, top of the upper mantle are collectively known as the
lithosphere, and it is of the lithosphere that the tectonic plates are comprised. Beneath the lithosphere is the
asthenosphere, a relatively low-viscosity layer on which the lithosphere rides. Important changes in crystal
structure within the mantle occur at 410 and 660 kilometers below the surface, spanning a transition zone
that separates the upper and lower mantle. Beneath the mantle, an extremely low viscosity liquid outer core
lies above a solid inner core.[68] The inner core may rotate at a slightly higher angular velocity than the
remainder of the planet, advancing by 0.1–0.5° per year.[69]
Geologic layers of the Earth[70]
Depth[ Density
Layer g/cm3

0–60 —
e 11]

0–35 Crust[note 12] 2.2–2.9

35–60 Upper mantle 3.4–4.4

Mantle 3.4–5.6

Asthenosphere —

Earth cutaway from core to exosphere. Not to 2890–

Outer core 9.9–12.2
scale. 5100

5100– 12.8–
Inner core
6378 13.1
[edit] Heat
Earth's internal heat comes from a combination of residual heat from planetary accretion (about 20%)
and heat produced through radioactive decay (80%).[72] The major heat-producing isotopes in the Earth are
potassium-40, uranium-238, uranium-235, and thorium-232.[73] At the center of the planet, the temperature
may be up to 7,000 K and the pressure could reach 360 GPa.[74] Because much of the heat is provided by
radioactive decay, scientists believe that early in Earth history, before isotopes with short half-lives had been
depleted, Earth's heat production would have been much higher. This extra heat production, twice present-
day at approximately 3 billion years ago,[72] would have increased temperature gradients within the Earth,
increasing the rates of mantle convection and plate tectonics, and allowing the production of igneous rocks
such as komatiites that are not formed today.[75]
Present-day major heat-producing isotopes[76]
Heat Half-life Mean mantle Heat
Isotope release concentration release
W/kg isotope years kg isotope/kg mantle W/kg mantle

238U 4.47 × 2.91 ×

9.46 × 10−5 30.8 × 10−9
109 10−12

235U 7.04 × 1.25 ×

5.69 × 10−4 0.22 × 10−9
108 10−13

232Th 1.40 × 3.27 ×

2.64 × 10−5 124 × 10−9
1010 10−12
40K 1.25 × 1.08 ×
2.92 × 10−5 36.9 × 10−9
109 10−12

Total heat loss from the Earth is 4.2 × 1013 watts.[77] A portion of the core's thermal energy is
transported toward the crust by mantle plumes; a form of convection consisting of upwellings of higher-
temperature rock. These plumes can produce hotspots and flood basalts.[78] More of the heat in the Earth is
lost through plate tectonics, by mantle upwelling associated with mid-ocean ridges. The final major mode of
heat loss is through conduction through the lithosphere, the majority of which occurs in the oceans because
the crust there is much thinner than that of the continents.[77]

[edit] Tectonic plates

Earth's main plates[79]
Plate name
106 km2

African Plate[note 13] 78.0

Antarctic Plate 60.9

Indo-Australian Plate 47.2

Eurasian Plate 67.8

North American Plate 75.9

South American Plate 43.6

Pacific Plate 103.3

Main article: Plate tectonics
The mechanically rigid outer layer of the Earth, the lithosphere, is broken into pieces called tectonic
plates. These plates are rigid segments that move in relation to one another at one of three types of plate
boundaries: Convergent boundaries, at which two plates come together, Divergent boundaries, at which two
plates are pulled apart, and Transform boundaries, in which two plates slide past one another laterally.
Earthquakes, volcanic activity, mountain-building, and oceanic trench formation can occur along these plate
boundaries.[80] The tectonic plates ride on top of the asthenosphere, the solid but less-viscous part of the
upper mantle that can flow and move along with the plates,[81] and their motion is strongly coupled with
convection patterns inside the Earth's mantle.
As the tectonic plates migrate across the planet, the ocean floor is subducted under the leading
edges of the plates at convergent boundaries. At the same time, the upwelling of mantle material at divergent
boundaries creates mid-ocean ridges. The combination of these processes continually recycles the oceanic
crust back into the mantle. Because of this recycling, most of the ocean floor is less than 100 million years in
age. The oldest oceanic crust is located in the Western Pacific, and has an estimated age of about 200
million years.[82][83] By comparison, the oldest dated continental crust is 4030 million years old.[84]
Other notable plates include the Indian Plate, the Arabian Plate, the Caribbean Plate, the Nazca
Plate off the west coast of South America and the Scotia Plate in the southern Atlantic Ocean. The Australian
Plate fused with the Indian Plate between 50 and 55 million years ago. The fastest-moving plates are the
oceanic plates, with the Cocos Plate advancing at a rate of 75 mm/yr[85] and the Pacific Plate moving 52–
69 mm/yr. At the other extreme, the slowest-moving plate is the Eurasian Plate, progressing at a typical rate
of about 21 mm/yr.[86]

[edit] Surface
Main articles: Landform and Extreme points of Earth
The Earth's terrain varies greatly from place to place. About 70.8%[87] of the surface is covered by
water, with much of the continental shelf below sea level. The submerged surface has mountainous features,
including a globe-spanning mid-ocean ridge system, as well as undersea volcanoes,[59] oceanic trenches,
submarine canyons, oceanic plateaus and abyssal plains. The remaining 29.2% not covered by water
consists of mountains, deserts, plains, plateaus, and other geomorphologies.
The planetary surface undergoes reshaping over geological time periods because of tectonics and
erosion. The surface features built up or deformed through plate tectonics are subject to steady weathering
from precipitation, thermal cycles, and chemical effects. Glaciation, coastal erosion, the build-up of coral
reefs, and large meteorite impacts[88] also act to reshape the landscape.

Present day Earth altimetry and bathymetry. Data from the National Geophysical Data Center's
TerrainBase Digital Terrain Model.
The continental crust consists of lower density material such as the igneous rocks granite and
andesite. Less common is basalt, a denser volcanic rock that is the primary constituent of the ocean floors.
[89] Sedimentary rock is formed from the accumulation of sediment that becomes compacted together.
Nearly 75% of the continental surfaces are covered by sedimentary rocks, although they form only about 5%
of the crust.[90] The third form of rock material found on Earth is metamorphic rock, which is created from the
transformation of pre-existing rock types through high pressures, high temperatures, or both. The most
abundant silicate minerals on the Earth's surface include quartz, the feldspars, amphibole, mica, pyroxene
and olivine.[91] Common carbonate minerals include calcite (found in limestone) and dolomite.[92]
The pedosphere is the outermost layer of the Earth that is composed of soil and subject to soil
formation processes. It exists at the interface of the lithosphere, atmosphere, hydrosphere and biosphere.
Currently the total arable land is 13.31% of the land surface, with only 4.71% supporting permanent crops.
[10] Close to 40% of the Earth's land surface is presently used for cropland and pasture, or an estimated
1.3 × 107 km2 of cropland and 3.4 × 107 km2 of pastureland.[93]
The elevation of the land surface of the Earth varies from the low point of −418 m at the Dead Sea, to
a 2005-estimated maximum altitude of 8,848 m at the top of Mount Everest. The mean height of land above
sea level is 840 m.[94]

[edit] Hydrosphere
Main article: Hydrosphere
Elevation histogram of the surface of the Earth
The abundance of water on Earth's surface is a unique feature that distinguishes the "Blue Planet"
from others in the Solar System. The Earth's hydrosphere consists chiefly of the oceans, but technically
includes all water surfaces in the world, including inland seas, lakes, rivers, and underground waters down to
a depth of 2,000 m. The deepest underwater location is Challenger Deep of the Mariana Trench in the Pacific
Ocean with a depth of −10,911.4 m.[note 14][95]
The mass of the oceans is approximately 1.35 × 1018 metric tons, or about 1/4400 of the total mass
of the Earth. The oceans cover an area of 361.8 × 106 km2 with a mean depth of 3,682 m, resulting in an
estimated volume of 1.332 × 109 km3.[96] If all the land on Earth were spread evenly, water would rise to an
altitude of more than 2.7 km.[note 15] About 97.5% of the water is saline, while the remaining 2.5% is fresh
water. Most fresh water, about 68.7%, is currently ice.[97]
The average salinity of the Earth's oceans is about 35 grams of salt per kilogram of sea water
(35 ‰).[98] Most of this salt was released from volcanic activity or extracted from cool, igneous rocks.[99]
The oceans are also a reservoir of dissolved atmospheric gases, which are essential for the survival of many
aquatic life forms.[100] Sea water has an important influence on the world's climate, with the oceans acting
as a large heat reservoir.[101] Shifts in the oceanic temperature distribution can cause significant weather
shifts, such as the El Niño-Southern Oscillation.[102]

[edit] Atmosphere
Main article: Atmosphere of Earth
The atmospheric pressure on the surface of the Earth averages 101.325 kPa, with a scale height of
about 8.5 km.[3] It is 78% nitrogen and 21% oxygen, with trace amounts of water vapor, carbon dioxide and
other gaseous molecules. The height of the troposphere varies with latitude, ranging between 8 km at the
poles to 17 km at the equator, with some variation resulting from weather and seasonal factors.[103]
Earth's biosphere has significantly altered its atmosphere. Oxygenic photosynthesis evolved 2.7
billion years ago, forming the primarily nitrogen-oxygen atmosphere of today. This change enabled the
proliferation of aerobic organisms as well as the formation of the ozone layer which blocks ultraviolet solar
radiation, permitting life on land. Other atmospheric functions important to life on Earth include transporting
water vapor, providing useful gases, causing small meteors to burn up before they strike the surface, and
moderating temperature.[104] This last phenomenon is known as the greenhouse effect: trace molecules
within the atmosphere serve to capture thermal energy emitted from the ground, thereby raising the average
temperature. Carbon dioxide, water vapor, methane and ozone are the primary greenhouse gases in the
Earth's atmosphere. Without this heat-retention effect, the average surface temperature would be −18 °C and
life would likely not exist.[87]

[edit] Weather and climate

Main articles: Weather and Climate

Satellite cloud cover image of Earth using NASA's Moderate-Resolution Imaging Spectroradiometer.
The Earth's atmosphere has no definite boundary, slowly becoming thinner and fading into outer
space. Three-quarters of the atmosphere's mass is contained within the first 11 km of the planet's surface.
This lowest layer is called the troposphere. Energy from the Sun heats this layer, and the surface below,
causing expansion of the air. This lower density air then rises, and is replaced by cooler, higher density air.
The result is atmospheric circulation that drives the weather and climate through redistribution of heat
The primary atmospheric circulation bands consist of the trade winds in the equatorial region below
30° latitude and the westerlies in the mid-latitudes between 30° and 60°.[106] Ocean currents are also
important factors in determining climate, particularly the thermohaline circulation that distributes heat energy
from the equatorial oceans to the polar regions.[107]

Source regions of global air masses

Water vapor generated through surface evaporation is transported by circulatory patterns in the
atmosphere. When atmospheric conditions permit an uplift of warm, humid air, this water condenses and
settles to the surface as precipitation.[105] Most of the water is then transported to lower elevations by river
systems and usually returned to the oceans or deposited into lakes. This water cycle is a vital mechanism for
supporting life on land, and is a primary factor in the erosion of surface features over geological periods.
Precipitation patterns vary widely, ranging from several meters of water per year to less than a millimeter.
Atmospheric circulation, topological features and temperature differences determine the average
precipitation that falls in each region.[108]
The amount of solar energy reaching the Earth's decreases with increasing latitude. At higher
latitudes the sunlight reaches the surface at a lower angles and it must pass through thicker columns of the
atmosphere. As a result, the mean annual air temperature at sea level decreases by about 0.4° C per per
degree of latitude away from the equator.[109] The Earth can be sub-divided into specific latitudinal belts of
approximately homogeneous climate. Ranging from the equator to the polar regions, these are the tropical
(or equatorial), subtropical, temperate and polar climates.[110] Climate can also be classified based on the
temperature and precipitation, with the climate regions characterized by fairly uniform air masses. The
commonly used Köppen climate classification system (as modified by Wladimir Köppen's student Rudolph
Geiger) has five broad groups (humid tropics, arid, humid middle latitudes, continental and cold polar), which
are further divided into more specific subtypes.[106]
[edit] Upper atmosphere

This view from orbit shows the full Moon partially obscured and deformed by the Earth's atmosphere.
NASA image.
See also: Outer space
Above the troposphere, the atmosphere is usually divided into the stratosphere, mesosphere, and
thermosphere.[104] Each layer has a different lapse rate, defining the rate of change in temperature with
height. Beyond these, the exosphere thins out into the magnetosphere, where the Earth's magnetic fields
interact with the solar wind.[111] An important part of the atmosphere for life on Earth is the ozone layer, a
component of the stratosphere that partially shields the surface from ultraviolet light. The Kármán line,
defined as 100 km above the Earth's surface, is a working definition for the boundary between atmosphere
and space.[112]
Thermal energy causes some of the molecules at the outer edge of the Earth's atmosphere have
their velocity increased to the point where they can escape from the planet's gravity. This results in a slow but
steady leakage of the atmosphere into space. Because unfixed hydrogen has a low molecular weight, it can
achieve escape velocity more readily and it leaks into outer space at a greater rate than other gasses.[113]
The leakage of hydrogen into space contributes to the pushing of the Earth from an initially reducing state to
its current oxidizing one. Photosynthesis provided a source of free oxygen, but the loss of reducing agents
such as hydrogen is believed to have been a necessary precondition for the widespread accumulation of
oxygen in the atmosphere.[114] Hence the ability of hydrogen to escape from the Earth's atmosphere may
have influenced the nature of life that developed on the planet.[115] In the current, oxygen-rich atmosphere
most hydrogen is converted into water before it has an opportunity to escape. Instead, most of the hydrogen
loss comes from the destruction of methane in the upper atmosphere.[116]
[edit] Magnetic field

The Earth's magnetic field, which approximates a dipole

Main article: Earth's magnetic field
The Earth's magnetic field is shaped roughly as a magnetic dipole, with the poles currently located
proximate to the planet's geographic poles. According to dynamo theory, the field is generated within the
molten outer core region where heat creates convection motions of conducting materials, generating electric
currents. These in turn produce the Earth's magnetic field. The convection movements in the core are
chaotic; the magnetic poles drift and periodically change alignment. This results in field reversals at irregular
intervals averaging a few times every million years. The most recent reversal occurred approximately
700,000 years ago.[117][118]
The field forms the magnetosphere, which deflects particles in the solar wind. The sunward edge of
the bow shock is located at about 13 times the radius of the Earth. The collision between the magnetic field
and the solar wind forms the Van Allen radiation belts, a pair of concentric, torus-shaped regions of energetic
charged particles. When the plasma enters the Earth's atmosphere at the magnetic poles, it forms the aurora.

[edit] Orbit and rotation

[edit] Rotation
Main article: Earth's rotation
Earth's axial tilt (or obliquity) and its relation to the rotation axis and plane of orbit.
Earth's rotation period relative to the Sun—its mean solar day—is 86,400 seconds of mean solar time.
Each second is slightly longer than an SI second because Earth's solar day is now slightly longer than it was
during the 19th century because of tidal acceleration.[120]
Earth's rotation period relative to the fixed stars, called its stellar day by the International Earth
Rotation and Reference Systems Service (IERS), is 86164.098903691 seconds of mean solar time (UT1), or
23h 56m 4.098903691s. [2][note 16] Earth's rotation period relative to the precessing or moving mean vernal
equinox, misnamed its sidereal day, is 86164.09053083288 seconds of mean solar time (UT1) (23 h 56m
4.09053083288s).[2] Thus the sidereal day is shorter than the stellar day by about 8.4 ms.[121] The length of
the mean solar day in SI seconds is available from the IERS for the periods 1623–2005[122] and 1962–2005.
Apart from meteors within the atmosphere and low-orbiting satellites, the main apparent motion of
celestial bodies in the Earth's sky is to the west at a rate of 15°/h = 15'/min. For bodies near the celestial
equator, this is equivalent to an apparent diameter of the Sun or Moon every two minutes; from the planet's
surface, the apparent sizes of the Sun and the Moon are approximately the same.[124][125]

[edit] Orbit
Main article: Earth's orbit
Earth orbits the Sun at an average distance of about 150 million kilometers every
365.2564 mean solar days, or one sidereal year. From Earth, this gives an apparent movement of the Sun
eastward with respect to the stars at a rate of about 1°/day, or a Sun or Moon diameter every 12 hours.
Because of this motion, on average it takes 24 hours—a solar day—for Earth to complete a full rotation about
its axis so that the Sun returns to the meridian. The orbital speed of the Earth averages about 30 km/s
(108,000 km/h), which is fast enough to cover the planet's diameter (about 12,600 km) in seven minutes, and
the distance to the Moon (384,000 km) in four hours.[3]
The Moon revolves with the Earth around a common barycenter every 27.32 days relative to the
background stars. When combined with the Earth–Moon system's common revolution around the Sun, the
period of the synodic month, from new moon to new moon, is 29.53 days. Viewed from the celestial north
pole, the motion of Earth, the Moon and their axial rotations are all counter-clockwise. Viewed from a vantage
point above the north poles of both the Sun and the Earth, the Earth appears to revolve in a counterclockwise
direction about the Sun. The orbital and axial planes are not precisely aligned: Earth's axis is tilted some
23.5 degrees from the perpendicular to the Earth–Sun plane, and the Earth–Moon plane is tilted about
5 degrees against the Earth-Sun plane. Without this tilt, there would be an eclipse every two weeks,
alternating between lunar eclipses and solar eclipses.[3][126]
The Hill sphere, or gravitational sphere of influence, of the Earth is about 1.5 Gm (or 1,500,000
kilometers) in radius.[127][note 17] This is maximum distance at which the Earth's gravitational influence is
stronger than the more distant Sun and planets. Objects must orbit the Earth within this radius, or they can
become unbound by the gravitational perturbation of the Sun.
Illustration of the Milky Way Galaxy, showing the location of the Sun
Earth, along with the Solar System, is situated in the Milky Way galaxy, orbiting about 28,000 light
years from the center of the galaxy. It is currently about 20 light years above the galaxy's equatorial plane in
the Orion spiral arm.[128]

[edit] Axial tilt and seasons

Main article: Axial tilt
Because of the axial tilt of the Earth, the amount of sunlight reaching any given point on the surface
varies over the course of the year. This results in seasonal change in climate, with summer in the northern
hemisphere occurring when the North Pole is pointing toward the Sun, and winter taking place when the pole
is pointed away. During the summer, the day lasts longer and the Sun climbs higher in the sky. In winter, the
climate becomes generally cooler and the days shorter. Above the Arctic Circle, an extreme case is reached
where there is no daylight at all for part of the year—a polar night. In the southern hemisphere the situation is
exactly reversed, with the South Pole oriented opposite the direction of the North Pole.

Earth and Moon from Mars, imaged by Mars Reconnaissance Orbiter. From space, the Earth can be
seen to go through phases similar to the phases of the Moon.
By astronomical convention, the four seasons are determined by the solstices—the point in the orbit of
maximum axial tilt toward or away from the Sun—and the equinoxes, when the direction of the tilt and the
direction to the Sun are perpendicular. In the northern hemisphere, Winter Solstice occurs on about
December 21, Summer Solstice is near June 21, Spring Equinox is around March 20 and Autumnal Equinox
is about September 23. In the Southern hemisphere, the situation is reversed, with the Summer and Winter
Solstices exchanged and the Spring and Autumnal Equinox dates switched.[129]
The angle of the Earth's tilt is relatively stable over long periods of time. However, the tilt does
undergo nutation; a slight, irregular motion with a main period of 18.6 years.[130] The orientation (rather than
the angle) of the Earth's axis also changes over time, precessing around in a complete circle over each
25,800 year cycle; this precession is the reason for the difference between a sidereal year and a tropical
year. Both of these motions are caused by the varying attraction of the Sun and Moon on the Earth's
equatorial bulge. From the perspective of the Earth, the poles also migrate a few meters across the surface.
This polar motion has multiple, cyclical components, which collectively are termed quasiperiodic motion. In
addition to an annual component to this motion, there is a 14-month cycle called the Chandler wobble. The
rotational velocity of the Earth also varies in a phenomenon known as length of day variation.[131]
In modern times, Earth's perihelion occurs around January 3, and the aphelion around July 4.
However, these dates change over time due to precession and other orbital factors, which follow cyclical
patterns known as Milankovitch cycles. The changing Earth-Sun distance results in an increase of about
6.9%[132] in solar energy reaching the Earth at perihelion relative to aphelion. Since the southern
hemisphere is tilted toward the Sun at about the same time that the Earth reaches the closest approach to
the Sun, the southern hemisphere receives slightly more energy from the Sun than does the northern over
the course of a year. However, this effect is much less significant than the total energy change due to the
axial tilt, and most of the excess energy is absorbed by the higher proportion of water in the southern

[edit] Moon
Diameter 3,474.8 km

Mass 7.349 × 1022 kg

Semi-major axis 384,400 km

Orbital period 27 d 7 h 43.7 m

Main article: Moon
The Moon is a relatively large, terrestrial, planet-like satellite, with a diameter about one-quarter of
the Earth's. It is the largest moon in the Solar System relative to the size of its planet, although Charon is
larger relative to the dwarf planet Pluto. The natural satellites orbiting other planets are called "moons" after
Earth's Moon.
The gravitational attraction between the Earth and Moon causes tides on Earth. The same effect on
the Moon has led to its tidal locking: its rotation period is the same as the time it takes to orbit the Earth. As a
result, it always presents the same face to the planet. As the Moon orbits Earth, different parts of its face are
illuminated by the Sun, leading to the lunar phases; the dark part of the face is separated from the light part
by the solar terminator.
Because of their tidal interaction, the Moon recedes from Earth at the rate of approximately 38 mm a
year. Over millions of years, these tiny modifications—and the lengthening of Earth's day by about 23 µs a
year—add up to significant changes.[134] During the Devonian period, for example, (approximately 410
million years ago) there were 400 days in a year, with each day lasting 21.8 hours.[135]

Details of the Earth-Moon system. Besides the radius of each object, the radius to the Earth-Moon
barycenter is shown. Photos from NASA. Data from NASA. The Moon's axis is located by Cassini's third law.
The Moon may have dramatically affected the development of life by moderating the planet's climate.
Paleontological evidence and computer simulations show that Earth's axial tilt is stabilized by tidal
interactions with the Moon.[136] Some theorists believe that without this stabilization against the torques
applied by the Sun and planets to the Earth's equatorial bulge, the rotational axis might be chaotically
unstable, exhibiting chaotic changes over millions of years, as appears to be the case for Mars.[137] If
Earth's axis of rotation were to approach the plane of the ecliptic, extremely severe weather could result from
the resulting extreme seasonal differences. One pole would be pointed directly toward the Sun during
summer and directly away during winter. Planetary scientists who have studied the effect claim that this
might kill all large animal and higher plant life.[138] However, this is a controversial subject, and further
studies of Mars—whose rotation period and axial tilt are similar to those of Earth, but which lacks a large
moon or liquid core—may settle the matter.
Viewed from Earth, the Moon is just far enough away to have very nearly the same apparent-sized
disk as the Sun. The angular size (or solid angle) of these two bodies match because, although the Sun's
diameter is about 400 times as large as the Moon's, it is also 400 times more distant.[125] This allows total
and annular solar eclipses to occur on Earth.
The most widely accepted theory of the Moon's origin, the giant impact theory, states that it formed
from the collision of a Mars-size protoplanet called Theia with the early Earth. This hypothesis explains
(among other things) the Moon's relative lack of iron and volatile elements, and the fact that its composition is
nearly identical to that of the Earth's crust.[139]
Earth has at least two co-orbital asteroids, 3753 Cruithne and 2002 AA29.[140]

A scale representation of the relative sizes of, and average distance between, Earth and Moon
[edit] Habitability
See also: Planetary habitability

A range of theoretical habitable zones with stars of different mass (our Solar System at center). Scale
is logarithmic, and planet sizes are not to scale.
A planet that can sustain life is termed habitable, even if life did not originate there. The Earth
provides the (currently understood) requisite conditions of liquid water, an environment where complex
organic molecules can assemble, and sufficient energy to sustain metabolism.[141] The distance of the Earth
from the Sun, as well as its orbital eccentricity, rate of rotation, axial tilt, geological history, sustaining
atmosphere and protective magnetic field all contribute to the conditions believed necessary to originate and
sustain life on this planet.[142]

[edit] Biosphere
Main article: Biosphere
The planet's life forms are sometimes said to form a "biosphere". This biosphere is generally believed
to have begun evolving about 3.5 billion years ago. Earth is the only place in the universe where life is known
to exist. Some scientists believe that Earth-like biospheres might be rare.[143]
The biosphere is divided into a number of biomes, inhabited by broadly similar plants and animals.
On land, biomes are separated primarily by differences in latitude, height above sea level and humidity.
Terrestrial biomes lying within the Arctic or Antarctic Circles, at high altitudes or in extremely arid areas are
relatively barren of plant and animal life; species diversity reaches a peak in humid lowlands at equatorial

[edit] Natural resources and land use

Main article: Natural resource
The Earth provides resources that are exploitable by humans for useful purposes. Some of these are
non-renewable resources, such as mineral fuels, that are difficult to replenish on a short time scale.
Large deposits of fossil fuels are obtained from the Earth's crust, consisting of coal, petroleum,
natural gas and methane clathrate. These deposits are used by humans both for energy production and as
feedstock for chemical production. Mineral ore bodies have also been formed in Earth's crust through a
process of Ore genesis, resulting from actions of erosion and plate tectonics.[145] These bodies form
concentrated sources for many metals and other useful elements.
The Earth's biosphere produces many useful biological products for humans, including (but far from
limited to) food, wood, pharmaceuticals, oxygen, and the recycling of many organic wastes. The land-based
ecosystem depends upon topsoil and fresh water, and the oceanic ecosystem depends upon dissolved
nutrients washed down from the land.[146] Humans also live on the land by using building materials to
construct shelters. In 1993, human use of land is approximately:
Arable Permanent Permanent Forests Urban
Land use O
land crops pastures and woodland areas

Percentage 4.71%[10] 26% 32% 1.5%

The estimated amount of irrigated land in 1993 was 2,481,250 km2.[10]

[edit] Natural and environmental hazards

Large areas are subject to extreme weather such as tropical cyclones, hurricanes, or typhoons that
dominate life in those areas. Many places are subject to earthquakes, landslides, tsunamis, volcanic
eruptions, tornadoes, sinkholes, blizzards, floods, droughts, and other calamities and disasters.
Many localized areas are subject to human-made pollution of the air and water, acid rain and toxic
substances, loss of vegetation (overgrazing, deforestation, desertification), loss of wildlife, species extinction,
soil degradation, soil depletion, erosion, and introduction of invasive species.
According to the United Nations, a scientific consensus exists linking human activities to global
warming due to industrial carbon dioxide emissions. This is predicted to produce changes such as the
melting of glaciers and ice sheets, more extreme temperature ranges, significant changes in weather and a
global rise in average sea levels.[147]

[edit] Human geography

Main article: Human geography
See also: World
Cartography, the study and practice of map making, and vicariously geography, have historically
been the disciplines devoted to depicting the Earth. Surveying, the determination of locations and distances,
and to a lesser extent navigation, the determination of position and direction, have developed alongside
cartography and geography, providing and suitably quantifying the requisite information.
Earth has approximately 6,803,000,000 human inhabitants as of December 12, 2009.[148]
Projections indicate that the world's human population will reach seven billion in 2013 and 9.2 billion in 2050.
[149] Most of the growth is expected to take place in developing nations. Human population density varies
widely around the world, but a majority live in Asia. By 2020, 60% of the world's population is expected to be
living in urban, rather than rural, areas.[150]
It is estimated that only one-eighth of the surface of the Earth is suitable for humans to live on—three-
quarters is covered by oceans, and half of the land area is either desert (14%),[151] high mountains (27%),
[152] or other less suitable terrain. The northernmost permanent settlement in the world is Alert, on
Ellesmere Island in Nunavut, Canada.[153] (82°28′N) The southernmost is the Amundsen-Scott South Pole
Station, in Antarctica, almost exactly at the South Pole. (90°S)
The Earth at night, a composite of DMSP/OLS ground illumination data on a simulated night-time
image of the world. This image is not photographic and many features are brighter than they would appear to
a direct observer.
Independent sovereign nations claim the planet's entire land surface, except for some parts of
Antarctica and the odd unclaimed area of Bir Tawil between Egypt and Sudan. As of 2007 there are 201
sovereign states, including the 192 United Nations member states. In addition, there are 59 dependent
territories, and a number of autonomous areas, territories under dispute and other entities.[10] Historically,
Earth has never had a sovereign government with authority over the entire globe, although a number of
nation-states have striven for world domination and failed.[154]
The United Nations is a worldwide intergovernmental organization that was created with the goal of
intervening in the disputes between nations, thereby avoiding armed conflict.[155] It is not, however, a world
government. The U.N. serves primarily as a forum for international diplomacy and international law. When
the consensus of the membership permits, it provides a mechanism for armed intervention.[156]
The first human to orbit the Earth was Yuri Gagarin on April 12, 1961.[157] In total, about 400 people
visited outer space and reached Earth orbit as of 2004, and, of these, twelve have walked on the Moon.[158]
[159][160] Normally the only humans in space are those on the International Space Station. The station's
crew, currently six people, is usually replaced every six months.[161] The furthest humans have travelled
from Earth is 400,171 km, achieved during the 1970 Apollo 13 mission.[162]
[edit] Cultural viewpoint

The first photograph ever taken by astronauts of an "Earthrise", from Apollo 8

Main article: Earth in culture
The name "Earth" derives from the Anglo-Saxon word erda, which means ground or soil, and is
related to the German word erde. It became eorthe later, and then erthe in Middle English.[163] The standard
astronomical symbol of the Earth consists of a cross circumscribed by a circle.[164]
Unlike the rest of the planets in the Solar System, mankind did not perceive the Earth as a planet
until the 16th century.[165] Earth has often been personified as a deity, in particular a goddess. In many
cultures the mother goddess is also portrayed as a fertility deity. Creation myths in many religions recall a
story involving the creation of the Earth by a supernatural deity or deities. A variety of religious groups, often
associated with fundamentalist branches of Protestantism[166] or Islam,[167] assert that their interpretations
of these creation myths in sacred texts are literal truth and should be considered alongside or replace
conventional scientific accounts of the formation of the Earth and the origin and development of life.[168]
Such assertions are opposed by the scientific community[169][170] and by other religious groups.[171][172]
[173] A prominent example is the creation-evolution controversy.
In the past there were varying levels of belief in a flat Earth,[174] but this was displaced by the
concept of a spherical Earth due to observation and circumnavigation.[175] The human perspective
regarding the Earth has changed following the advent of spaceflight, and the biosphere is now widely viewed
from a globally integrated perspective.[176][177] This is reflected in a growing environmental movement that
is concerned about humankind's effects on the planet.[178]

[edit] See also

Solar System portal

Earth sciences portal

Book:Solar System

Books are collections of articles that can be downloaded or ordered in print.

• List of Earth-related topics
• Earth science
• Outline of earth science
• Index of earth science articles
• Geodesy
• History of geodesy
• Geography
• List of basic geography topics
• Index of geography articles
• Geology
• Outline of geology
• Index of geology articles

[edit] Notes
1. ^ All astronomical quantities vary, both secularly and periodically. The quantities given are
the values at the instant J2000.0 of the secular variation, ignoring all periodic variations.
2. ^ a b aphelion = a × (1 + e); perihelion = a × (1 - e), where a is the semi-major axis and e is
the eccentricity.
3. ^ The reference lists the longitude of the ascending node as -11.26064°, which is equivalent
to 348.73936° by the fact that any angle is equal to itself plus 360°.
4. ^ The reference lists the longitude of perihelion, which is the sum of the longitude of the
ascending node and the argument of perihelion. That is, 114.20783° + (-11.26064°) = 102.94719°.
5. ^ Due to natural fluctuations, ambiguities surrounding ice shelves, and mapping conventions
for vertical datums, exact values for land and ocean coverage are not meaningful. Based on data
from the Vector Map and Global Landcover datasets, extreme values for coverage of lakes and
streams are 0.6% and 1.0% of the earth's surface. The ice shields of Antarctica and Greenland are
counted as land, even though much of the rock which supports them lies below sea level.
6. ^ Blue Planet is used as the title of several films Blue Planet and The Blue Planet, in the Life
issue The Incredible Year '68 featuring the Earthrise photo with lines from poet James Dickey
Behold/The blue planet steeped in its dream/Of reality , and in the title of the European Space Agency
bulletin report Exploring the water cycle of the 'Blue Planet'
7. ^ By International Astronomical Union convention, the term terra is used only for naming
extensive land masses on celestial bodies other than the Earth. Cf. Blue, Jennifer (2007-07-05).
"Descriptor Terms (Feature Types)". Gazetteer of Planetary Nomenclature. USGS. Retrieved 2007-07-05.
8. ^ Other planets in the Solar System are either too hot or too cold to support liquid water.
However, it is confirmed to have existed on the surface of Mars in the past, and may still appear
today. See:
• Malik, Tariq (2007-03-02). "Rover reveals Mars was once wet enough for life". (via MSNBC). Retrieved 2007-08-28.
• Staff (2005-11-07). "Simulations Show Liquid Water Could Exist on Mars". Daily
Headlines (University of Arkansas). Retrieved 2007-
9. ^ As of 2007, water vapor has been detected in the atmosphere of only one extrasolar planet,
and it is a gas giant. See: Tinetti, G.; Vidal-Madjar, A.; Liang, M.C.; Beaulieu, J. P.; Yung, Y.; Carey,
S.; Barber, R. J.; Tennyson, J.; Ribas, I (July 2007). "Water vapour in the atmosphere of a transiting
extrasolar planet". Nature 448 (7150): 169–171. doi:10.1038/nature06002. PMID 17625559.
10.^ The number of solar days is one less than the number of sidereal days because the orbital
motion of the Earth about the Sun results in one additional revolution of the planet about its axis.
11.^ Locally varies between 5 and 200 km.
12.^ Locally varies between 5 and 70 km.
13.^ Including the Somali Plate, which is currently in the process of formation out of the African
Plate. See: Chorowicz, Jean (October 2005). "The East African rift system". Journal of African Earth
Sciences 43 (1–3): 379–410. doi:10.1016/j.jafrearsci.2005.07.019.
14.^ This is the measurement taken by the vessel Kaikō in March 1995 and is believed to be the
most accurate measurement to date. See the Challenger Deep article for more details.
15.^ The total surface area of the Earth is 5.1 × 108 km2. To first approximation, the average
depth would be the ratio of the two, or 2.7 km.
16.^ Aoki, the ultimate source of these figures, uses the term "seconds of UT1" instead of
"seconds of mean solar time".—Aoki, S. (1982). "The new definition of universal time". Astronomy and
Astrophysics 105 (2): 359–361. Retrieved 2008-
17.^ For the Earth, the Hill radius is
where m is the mass of the Earth, a is an Astronomical Unit, and M is the mass of the Sun. So the

radius in A.U. is about: .

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[edit] Further reading

• Comins, Neil F. (2001). Discovering the Essential Universe (Second ed.). W. H. Freeman.
ISBN 0-7167-5804-0. Retrieved 2007-03-17.
[edit] External links
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• The Gateway to Astronaut Photography of Earth
• "Global Measured Extremes of Temperature and Precipitation" . National Climatic Data
Center. August 20, 2008. Retrieved 2010-

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Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir Terre (homonymie).

La Terre vue depuis Apollo 17 en 1972[Note 1].
Caractéristiques orbitales
(Époque J2000.0)

Demi-grand axe 149 597 887,5 km

(1,0000001124 UA)

152 097 701 km

(1,0167103335 UA)

147 098 074 km

(0,9832898912 UA)

924 375 700 km

Circonférence orbitale
(6,1790699007 UA)

Excentricité 0,01671022

Période de révolution 365,25696 d

Période synodique —d

Vitesse orbitale moyenne 29,783 km/s

Vitesse orbitale maximale 30,287 km/s

Vitesse orbitale minimale 29,291 km/s

Inclinaison sur l'écliptique (par définition) 0°

Nœud ascendant 174,873°

Argument du périhélie 288,064°

Satellites connus 1, la Lune

Caractéristiques physiques

Rayon équatorial 6 378,137 km

Rayon polaire 6 356,752 km

Rayon moyen
6 371,0 km

Aplatissement 0,0033529

40 075,017 ;
Périmètre équatorial
périmètre méridional "polaire" = 40 007,864 km

Superficie 510 067 420 km2

Volume 1,08321×1012 km3

Masse 5,9736×1024 kg

Masse volumique globale 5,515×103 kg/m3

Gravité de surface 9,780 m/s2
(0,99732 g)

Vitesse de libération 11,186 km/s

Période de rotation 0,99726949 d

(jour sidéral) (23 h 56 min 4,084 s)

Vitesse de rotation
1 674,364 km/h
(à l’équateur)

Inclinaison de l’axe 23,4388°

Albédo géométrique visuel 0,367

Albédo de Bond 0,306

Irradiance solaire 1 367,6 W/m2

(1 Terre)

Température d'équilibre
254,3 K (-18,7 °C)
du corps noir

Température de surface :

Maximum : 57,8 °C[1]

Moyenne : 15 °C

Minimum : -89,2 °C[2]

Caractéristiques de l’atmosphère

Pression atmosphérique 101 325 Pa

Masse volumique au sol 1,217 kg/m3

Masse totale 5,1×1018 kg

Hauteur 8,5 km

Masse molaire moyenne 28,97 g/mol

Azote N2 78,084 % volume sec

Oxygène O2 20,946 % volume sec

Argon Ar 0,9340 % volume sec

Dioxyde de carbone CO2 390 ppm volume sec

Néon Ne 18,18 ppm volume sec

Hélium He 5,24 ppm volume sec

Méthane CH4 1,79 ppm volume sec

Krypton Kr 1,14 ppm volume sec

Hydrogène H2 550 ppb volume sec

Protoxyde d'azote N2O 300 ppb volume sec

Monoxyde de carbone CO 100 ppb volume sec

Xénon Xe 90 ppb volume sec

Ozone O3 0 à 70 ppb volume sec

Dioxyde d'azote NO2 20 ppb volume sec

Iode I 10 ppb volume sec

Vapeur d'eau H2O ~ 0,4 % volume global

~ de 1 à 4 % en surface (valeurs typiques)

• Nature planétaire pressentie par

l'école pythagoricienne (Philolaos de Crotone).
Découverte par
• Attestée à
l'époque hellénistique (Aristarque de Samos, puis Ératosthène).

• Ve siècle av. J.-C.

Découverte le
• IIIe siècle av. J.-C.
La Terre est la troisième planète du Système solaire par ordre de distance croissante au Soleil, et la
quatrième par taille et par masse croissantes. C'est la plus grande et la plus massive des quatre planètes
telluriques, les trois autres étant Mercure, Vénus et Mars.
Elle possède un satellite naturel, la Lune, qui est le cinquième plus gros satellite du Système solaire.
Elle est dotée d'un puissant champ magnétique qui dirige vers les pôles les particules chargées
véhiculées par le vent solaire, y provoquant des aurores polaires et générant des ceintures de radiations
concentriques autour du globe résultant de l'accumulation de ces particules piégées dans le champ
magnétique de la Terre. La magnétosphère agit ainsi comme un bouclier protégeant notre planète du vent
Couramment appelée en français Terre, planète Terre, planète bleue ou encore Monde[Note 2], c'est
une planète à manteau actif, dotée d'une atmosphère comportant de l'oxygène, recouverte d'eau liquide.
• 1 Histoire
• 2 Composition et structure
• 2.1 Composition chimique
• 2.2 Structure géologique
• 2.3 Plaques tectoniques
• 3 Atmosphère
• 3.1 Constitution
• 3.2 Structure de l'atmosphère
• 4 Données physiques et astronomiques
• 4.1 Forme de la Terre
• 4.2 Données orbitales
• 4.3 Jour, saisons et année
• 4.4 Accélération de la pesanteur
• 4.5 Satellites de la Terre
• 4.6 Position dans l'Univers
• 5 Jour de la Terre
• 6 Notes
• 7 Références
• 8 Voir aussi
Histoire [modifier]
Article détaillé : Histoire de la Terre.
La Terre ainsi que les autres planètes du système solaire se sont formées il y a 4,467 milliards
d'années à partir d'une nébuleuse solaire, masse de poussières et de gaz en forme de disque détachée du
Soleil en formation. Initialement en fusion, la couche externe de la Terre s'est refroidie pour former une
croûte solide. Plus tard, les impacts d'astéroïdes ont causé de nombreux changements sur l'environnement à
la surface. La Lune s'est formée peu de temps après, sans doute à la suite d'une collision avec un objet de la
taille de Mars (quelquefois appelé Théia). Une partie de cet objet se serait agglomérée avec la Terre, tandis
qu'une autre portion, mêlée avec peut-être 10 % de la masse totale de la Terre, aurait été éjectée dans
l'espace, où elle aurait formé la Lune.
L'activité volcanique a produit une atmosphère primitive. De la vapeur d'eau condensée, mêlée à de
la glace apportée par des comètes, a produit les océans. On suppose qu'une activité chimique intense dans
un milieu hautement énergétique a produit une molécule capable de se reproduire, il y a environ 4 milliards
d'années. La vie elle-même serait apparue 500 000 ans plus tard.
L'apparition de la photosynthèse met ensuite l'énergie solaire au service de la vie. Il en résulte en
effet à la fois une accumulation de dioxygène dans l'atmosphère, favorisant la vie animale, et le
développement d'une couche d'ozone[Note 3] dans la haute atmosphère, protégeant la surface de la planète
de l'agression des rayons ultraviolets. Dans ce nouveau cadre, la vie évolue de plus en plus vite vers des
formes toujours plus complexes.
La surface du globe se transforme continuellement, sur des périodes de plusieurs centaines de
millions d'années. Des continents ou supercontinents se forment puis se divisent. C'est ainsi qu'il y a environ
750 millions d'années, le plus vieux des supercontinents connus, Rodinia, commença à se disloquer. Les
continents entre lesquels il s'était divisé se recombinèrent plus tard pour former Pannotia, il y a 650-540
millions d'années, puis finalement Pangée, au Permien, qui se fragmenta il y a 180 millions d'années.
Depuis les années 1960, de nombreuses hypothèses ont été émises dont une qui affirme qu'une (ou
une série) de grande(s) glaciation(s) eut lieu il y a 750 et 580 millions d'années, pendant le
Néoprotérozoïque, et qui couvrit la planète d'une couche de glace. Cette hypothèse a été nommée Snowball
Earth (« Terre boule de neige »), et est d'un intérêt particulier parce qu'elle précède l'explosion cambrienne,
quand des formes de vies multicellulaires commencèrent à proliférer.
À la suite de l'explosion cambrienne, il y a 535 millions d'années, 5 extinctions massives eurent lieu.
La dernière extinction majeure date de 65 millions d'années, quand une présumée météorite est entrée en
collision avec la Terre, exterminant les dinosaures et d'autres grands reptiles, épargnant de plus petits
animaux comme les mammifères, oiseaux, lézards, etc. Dans les 65 millions d'années qui se sont écoulées
depuis, les mammifères se sont diversifiés, l'espèce humaine s'étant développée depuis deux millions
d'années. Des changements périodiques à long terme de l'orbite de la Terre, causés par l'influence
gravitationnelle des autres astres, sont probablement une des causes des glaciations qui ont couvert une
bonne partie de la planète. À l'issue de la dernière glaciation, le développement de l'agriculture et, ensuite,
des civilisations, permit aux humains de modifier la surface de la Terre dans une courte période de temps,
comme aucune autre espèce avant lui sur terre, affectant la nature tout comme les autres formes de vies.

Composition et structure [modifier]

La Terre est une planète tellurique, c'est-à-dire une planète essentiellement rocheuse à noyau
métallique, contrairement aux géantes gazeuses, telles que Jupiter, essentiellement constituées de gaz
légers (hydrogène et hélium).
Il s'agit de la plus grande des quatre planètes telluriques du système solaire, que ce soit en termes
de taille ou de masse. De ces quatre planètes, la Terre a aussi la masse volumique globale la plus élevée, la
plus forte gravité de surface et le plus puissant champ magnétique global. Cependant, plusieurs planètes
telluriques plus grandes que la Terre ont été découvertes en dehors du système solaire, parmi lesquelles
l'exoplanète Gliese 581 c, qui possède un diamètre 50 % supérieur à celui de la Terre. Plusieurs missions
sont en cours, ou prévues, afin de découvrir de nouvelles planètes similaires à la Terre, appelées exoterres.
La surface externe de la Terre est divisée en plusieurs segments rigides, ou plaques tectoniques, qui
se déplacent lentement sur la surface sur une durée de plusieurs millions d'années. Environ 71 % de la
surface est couverte d'océans d'eau salée, les 29 % restants consistant en continents et îles. L'eau liquide,
nécessaire à la vie telle que nous la connaissons, est très abondante sur Terre, et aucune autre planète n'a
encore été découverte avec des étendues d'eau liquide (lacs, mers, océans) à sa surface.

Composition chimique [modifier]

La masse de la Terre est d'approximativement 5,98×1024 kg. Elle est composée principalement de
fer (32,1 %[3]), d'oxygène (30,1 %), de silicium (15,1 %), de magnésium (19,9 %), de soufre (2,9 %), de
nickel (1,8 %), de calcium (1,5 %) et d'aluminium (1,4 %), le 1,2 % restant consistant en de légères traces
d'autres éléments. Les éléments les plus denses ayant tendance à se concentrer au centre de la Terre
(phénomène de différenciation planétaire), on pense que le cœur de la Terre est composé majoritairement
de fer (88,8 %), avec une plus petite quantité de nickel (5,8 %), de soufre (4,5 %) et moins de 1 % d'autres
Le géochimiste F. W. Clarke a calculé que 47 % (en poids) de la croûte terrestre est faite d'oxygène,
présent principalement sous forme d'oxydes, dont les principaux sont les oxydes de silicium, aluminium, fer,
calcium, magnésium, potassium et sodium. La silice est le constituant majeur de la croûte, sous forme de
pyroxénoïdes, les minéraux les plus communs des roches magmatiques et métamorphiques. Après une
synthèse basée sur l'analyse de 1 672 types de roches, Clarke a obtenu les pourcentages présentés dans le
tableau ci-dessous.

Silice (SiO2) 59,71

Oxyde d'aluminium (Al2O3) 15,41

Oxyde de calcium (CaO) 4,90

Oxyde de magnésium (MgO) 4,36

Oxyde de sodium (Na2O) 3,55

Oxyde de fer(II) (FeO) 3,52

Oxyde de potassium (K2O) 2,80

Oxyde de fer(III) (Fe2O3) 2,63

Eau (H2O) 1,52

Dioxyde de titane (TiO2) 0,60

Pentoxyde de phosphore (P2O5) 0,22

Total 99,22

Structure géologique [modifier]

Structure de la Terre.
1. croûte continentale,
2. croûte océanique,
3. manteau supérieur,
4. manteau inférieur,
5. noyau externe,
6. noyau interne,
A : Discontinuité de Mohorovicic,
B : Discontinuité de Gutenberg,
C : Discontinuité de Lehmann.
Article détaillé : Structure interne de la Terre.
La Terre est constituée de plusieurs couches internes identifiables à peu près concentriques : la
croûte terrestre (océanique ou continentale), le manteau supérieur, le manteau inférieur, et le noyau externe
et interne. La lithosphère est constituée de la croûte et de la zone superficielle du manteau supérieur.
L'asthénosphère est la zone plus profonde du manteau supérieur (en dessous de la lithosphère).
La croûte terrestre est relativement jeune par rapport à la Terre elle-même. Pendant la période
relativement courte d'environ 500 millions d'années pendant laquelle l'érosion et les processus tectoniques
ont détruit, puis recréé, la plupart des couches superficielles de la Terre, la presque totalité des traces de
l'histoire géologique de sa surface (cratères d'impact, par exemple) ont disparu.
Plus de 99 % de la surface terrestre aurait moins de 2 milliards d'années.
La structure interne de la Terre est connue au moyen de l'étude de la propagation des ondes
sismiques entre une source et différents points de la surface terrestre.
La vitesse d'une onde sismique change en effet assez brutalement au passage entre deux couches
de composition ou phase minérale différentes. Ces limites ont parfois reçu des noms particuliers, tels que la
discontinuité de Mohorovicic, la discontinuité de Lehmann ou la discontinuité de Gutenberg.
La constitution de la Terre s'explique par son mode de formation, par accrétion de météorites, qui a
produit une stratification en phase fluide par masse volumique décroissante depuis les couches internes vers
les couches externes.
La plus grande partie de la chaleur interne de la Terre (87 %) est produite par la radioactivité des
roches qui constituent la croûte terrestre : radioactivité naturelle produite par la désintégration de l'uranium,
du thorium et du potassium.

Plaques tectoniques [modifier]

Selon la théorie de la tectonique des plaques, la partie supérieure de l'intérieur de la Terre est
composée de deux couches : la lithosphère, comprenant la croûte, et la partie solide du manteau. Au-
dessous de la lithosphère se trouve l'asthénosphère, qui forme le cœur du manteau. L'asthénosphère
ressemble à du liquide extrêmement chaud et visqueux.
La lithosphère flotte essentiellement sur l'asthénosphère et est brisée en pièces qui sont appelées
plaques tectoniques. Ces plaques sont des segments rigides qui bougent en relation avec les autres de trois
façons : en convergence, en divergence, et par transcurrence. C'est ainsi que sont créés les tremblements
de terre, l'activité volcanique ainsi que les montagnes.
Certaines plaques ont une plus petite superficie comme la plaque indienne, la plaque arabique, la
plaque caraïbe et la plaque de Nazca à l'ouest de la côte de l'Amérique du Sud. La plaque australienne s'est
fusionnée quelque peu à la plaque indienne il y a 50 à 55 millions d'années. Les plaques les plus rapides
dans leur mouvement sont les plaques océaniques, se déplaçant d'environ 70 Millimètresmm/an. À l'opposé,
la plaque la plus lente est la plaque eurasienne, progressant d'environ 21 Millimètresmm/an.
Les principales plaques tectoniques sont :
Nom de la
Carte totale, en (106 Couvre

Plaque africaine 61,3 Afrique

Carte des plaques tectoniques terrestres. Les
flèches indiquent les mouvements relatifs de chaque Plaque
60,9 Antarctique
plaque. antarctique

47,2 Australie

Plaque Asie et
eurasienne l'Europe

Plaque nord- 75,9 Amérique

du Nord et Nord-Est
de la Sibérie

Plaque sud- Amérique

américaine du Sud

Plaque pacifique 103,3
Atmosphère [modifier]
Schéma des couches de l'atmosphère.
La Terre est entourée d'une enveloppe gazeuse qu'elle retient par attraction gravitationnelle :
l'atmosphère. L'atmosphère de la Terre est intermédiaire entre celle, très épaisse, de Vénus, et celle, très
ténue, de Mars : sa pression au niveau de la mer est en moyenne de 101 325 Pa, soit 1 atm par définition.
Outre une proportion variable de vapeur d'eau comprise entre 0 et 4 %, elle est constituée de 78,09 %
d'azote, 20,95 % d'oxygène,0,93 % d'argon et 0,039 %9 % de dioxyde de carbone, ainsi que de divers autres
gaz. Ce taux élevé d'oxygène est unique dans le Système solaire, et résulte de l'activité photosynthétique
des organismes chlorophylliens : la Terre est, en effet, le seul astre connu pour abriter la vie, conséquence
probable du fait que c'est également le seul astre connu pour avoir des conditions de température et de
pression permettant l'existence d'eau liquide en surface.
Cette atmosphère donne à la planète un reflet bleuté depuis l'espace, d'où son surnom de « planète
bleue ». La constitution et la densité de l'atmosphère sont telles que la lumière incidente du Soleil et la
lumière réfléchie par les continents et les mers sont diffractées ; donnant sa couleur au ciel, et par réflexion,
aux étendues d'eau.

Constitution [modifier]
Cette enveloppe, dont la masse globale est de l'ordre de 5×10 18 kg (un millionième de la masse de
la Terre), est contenue à 99 % dans les 30 premiers kilomètres (50 % dans les 5 premiers kilomètres).
La basse atmosphère (du niveau de la mer jusqu'à environ 45 km) est composée de gaz
« permanents », gaz dont les proportions restent constantes, et de gaz de concentration variable avec
• Le diazote, le dioxygène et l'argon constituent, en volume, 99,997 % des gaz permanents
(voir tableau ci-dessus) ; le brassage vertical de l'air permet de conserver une répartition constante à
tous les niveaux, même pour les gaz les plus légers, tels que l'hélium ou l'hydrogène.
• Les gaz à concentration variable sont essentiellement la vapeur d'eau H2O ; et dans une
moindre mesure le dioxyde de carbone CO2, le dioxyde de soufre SO2 et l'ozone O3.

L'atmosphère terrestre peut être considérée, à un instant donné, comme un mélange

thermodynamique d'air sec et de vapeur d'eau.
Les particules liquides, solides, ou mixtes, en suspension dans l'atmosphère constituent l'aérosol
Ces particules jouent un rôle primordial dans les phénomènes de condensation (nuages) et de
formation de cristaux de glace, ainsi qu'à différents processus physico-chimiques dans l'atmosphère. Leur
concentration varie de plusieurs puissances de 10 (de plusieurs ordres de grandeurs) en fonction du lieu et
du temps ; en concentration élevée, elles constituent un facteur de pollution. Les particules se classent en :
• particules d'Aitken : 1 nm < d < 0,1 µm
• grosses particules : 0,1 µm < d < 5 µm
• particules géantes : 5 µm < d < 50 µm environ
L'atmosphère atténue de façon importante le rayonnement solaire reçu au sol ; suivant l'importance
de la couverture nuageuse, le sol reçoit de 68 % à 28 % (ou moins) du rayonnement solaire parvenant à
l'atmosphère, un flux solaire initial de 1 370 W/m2.
Structure de l'atmosphère [modifier]
La composition chimique de l'atmosphère, sa température, ou les phénomènes qui y sont observés
présentent des discontinuités marquées lorsque l'altitude augmente. Ces discontinuités correspondent à des
couches homogènes dont les propriétés évoluent de façon continue ; ce sont (par altitude croissante) :
• la troposphère
• la stratosphère
• la mésosphère
• la thermosphère
• l'exosphère
Les limites de ces couches (d'altitude variable) ont reçu des désignations particulières : tropopause,
stratopause, mésopause et thermopause.

Données physiques et astronomiques [modifier]

Forme de la Terre [modifier]

La forme de la Terre est modélisée par un ellipsoïde, légèrement aplati aux pôles, et plus
précisément par le géoïde. Le diamètre approximatif de référence est de 12 742 km.
La rotation de la Terre crée un léger bourrelet équatorial, de sorte que le diamètre à l’équateur est
43 km plus long que le diamètre polaire (du pôle Nord au pôle Sud). Les plus grandes dénivellations du sol
de la Terre sont l'Everest (8 848 m au-dessus du niveau de la mer) et la fosse des Mariannes (11 500 m
sous le niveau de la mer). Par contre, à cause de l’aplatissement, l’objet le plus éloigné du cœur de la Terre
est en fait le volcan Chimborazo en Équateur.

Comparaison de quelques caractéristiques physiques des quatre planètes telluriques

Planète Rayon équatorial Masse Gravité

2 439,7 km (0,383 e23/3.3023,302×102

Mercure[4] 3,701 m/s² (0,377 g) ~
Terre) 3 kg (0,055 Terre)

Vénus[5] 6 051,8 km (0,95 Terre) 8,87 m/s² (0,904 g) 1
024 kg (0,815 Terre)

e24/5.97365,9736×1 9,780 m/s² (0,99732

Terre[6] 6 378,14 km 2
024 kg g)

3 402,45 km (0,533 e23/6.41856,4185×1

Mars[7] 3,69 m/s² (0,376 g) 2
Terre) 023 kg (0,107 Terre)
Photomontage comparatif des tailles des planètes telluriques (de gauche à droite) : Mercure, Vénus, la Terre et Ma

La conception sphérique de la Terre remonte à l'antiquité grecque, vers le Ve siècle av. J.-C., et plus
spécifiquement aux pythagoriciens. On retrouve cette conception chez Parménide, Platon ou Aristote. Elle
s'appuie sur le fait que, lors des éclipses de Lune, l'ombre de la Terre est sphérique, et sur le fait que les
constellations varient lorsqu'on se déplace du Nord au Sud[8]. Au IIIe siècle av. J.-C., Ératosthène donna une
estimation du rayon terrestre que nous supposons[Note 5] très proche de la réalité, ainsi que Posidonios, au
Ie siècle av. J.-C.. Vers la même époque, Cléomède définit les notions d'équateur, de tropiques, d'arctique et
d'antarctique. Reprenant ces notions, le géographe Ptolémée fournit au IIe siècle des informations
géographiques qui furent utilisées jusqu'à la Renaissance.
La civilisation arabo-musulmane conserva la connaissance d'une Terre sphérique et au IXe siècle, le
calife Al-Mamun, à Bagdad, fit procéder à une mesure d'une partie d'un méridien, conduisant à une bonne
approximation de la circonférence de la Terre. Plus explicite encore est qu'une figure comme Abou Hanîfa,
fondateur de l'une des quatre écoles de jurisprudence musulmane, avait foi dans la sphéricité de la Terre[9].
Dans le monde chrétien, cette idée fut parfois remise en cause, par exemple au VIe siècle par Cosmas
Indicopleustès. En effet, les Pères de l'Église ne pouvaient concilier la vision du monde sphérique d'Aristote
constitué de deux zones polaires et deux zones tempérées, séparées par une zone torride infranchissable,
avec l'universalité du message du Christ, ce message ne pouvant parvenir à d'hypothétiques[Note 6]
habitants des antipodes. Jusqu'au XIIe siècle, on s'attacha donc à représenter le monde sous forme
symbolique, mais des philosophes ou des religieux tels Isidore de Séville, Bède le Vénérable, Jean Scot
Erigène, Gerbert d'Aurillac, Thomas d'Aquin, Albert le Grand ou Roger Bacon avaient très bien intégré la
représentation sphérique. Vers 1150, le livre De imagine mundi, dont l'auteur présumé est Honorius d'Autun,
décrit l'univers de façon plus mythologique que scientifique, mais dans lequel la Terre est une sphère
d'environ 35 000 km de circonférence. Charlemagne est d'ailleurs représenté sur quelques enluminures
comme tenant à la main une représentation d'un petit globe terrestre surmonté d'une croix.
Position et taille du continent asiatique selon Christophe Colomb.
Les récits de voyages de missionnaires, de Marco Polo et de l'explorateur Jean de Mandeville (avec
son Livre des merveilles du monde) diffusaient dans la société l'image d'une terre sphérique, qui pouvait
théoriquement faire l'objet d'une « circumnavigation ». L’Imago mundi du cardinal Pierre d'Ailly retenait cette
représentation sphérique. On sait que Christophe Colomb a été influencé par le Livre des merveilles du
monde de Mandeville, et qu'il possédait un exemplaire de l’Imago mundi abondamment annoté et commenté
par ses soins. En sous-estimant grandement le rayon terrestre et en imaginant un continent asiatique trois
fois plus étendu vers l'Est qu'il ne l'est en réalité, Colomb a pu envisager de façon raisonnable la possibilité
de rejoindre les Indes par l'Ouest. Une connaissance plus précise des distances aurait découragé toute
tentative de traversée de l'Océan avec les moyens de l'époque.
Les voyages des Portugais dès le début du XVe siècle pour rejoindre les Indes en contournant
l'Afrique, la redécouverte des textes grecs à la Renaissance, en particulier la Géographie de Ptolémée, leur
diffusion au moyen de l'imprimerie ont également largement contribué à propager les représentations
modernes de la Terre, avec le Nord vers le haut des cartes, les méridiens, les parallèles, l'équateur et les
deux tropiques. Le plus ancien globe terrestre connu est fabriqué par Martin Behaim vers la fin du
XVe siècle, peu avant que Vasco de Gama, Christophe Colomb ou Magellan entreprennent leurs voyages.
On y voit l'Europe, l'Afrique et l'Asie, mais bien entendu, ni les Amériques, ni l'Océanie.
Mercator a, en dessinant ses cartes, mentionné et dessiné un énorme continent austral : Terra
incognita australis (terre australe (du sud) inconnue). Cette « terre australe inconnue » a été dessinée au
Sud car Mercator pensait, à la suite des Grecs, que sans ce poids la Terre n'était pas équilibrée. Les
réflexions et travaux en géographie (relevés cartographiques, projection de Mercator) au XVIe siècle ont
permis de faire évoluer la connaissance de la Terre.
C'est au XVIIIe siècle que l'aplatissement des pôles est reconnu, avec les expéditions menées au
Pérou et en Laponie.

Données orbitales [modifier]

Le demi-grand axe de son orbite définit l'unité astronomique, qui vaut 149 597 870 700 ± 3 m selon
la valeur actuellement la plus précise[10]. Son éloignement au Soleil est compris entre 0,9833 et 1,0167 UA.

Jour, saisons et année [modifier]

Article connexe : Année (astronomie).
La Terre tourne autour d'elle même en un jour sidéral de 23h56', et autour du soleil en une année
d'environ 365 jours. Son axe de rotation est incliné par rapport à l'écliptique, ce qui produit l'alternance de
quatre saisons sur la surface du globe.
La périodicité des saisons se mesure, elle, au moyen de l'année tropique, légèrement plus courte
que l'année sidérale de la valeur annuelle de la précession de l'axe de rotation. La période du passage aux
points de l'ellipse de l'orbite comme le périhélie, est l'année anomalistique, légèrement plus longue que
l'année sidérale de la valeur annuelle de la récession du périhélie.
Enfin, dans les calculs astronomiques, on utilise l'année julienne valant exactement 365,25 jours.

Accélération de la pesanteur [modifier]

Pour consulter un article plus général, voir : Pesanteur.
L'accélération de la pesanteur (ou « champ de pesanteur ») varie légèrement à la surface de la Terre
pour trois raisons :
• Elle dépend de l'altitude, l'accélération étant inversement proportionnelle au carré de la
distance entre le centre de gravité de la Terre et le point où il est mesuré.
• La Terre n'est pas parfaitement sphérique, mais un peu aplatie aux pôles, la gravitation est
plus grande aux pôles, pour la même raison.
• La Terre tourne sur elle-même, ce qui fait qu'un objet à l'équateur est un tout petit peu plus
léger (voir Force centrifuge).
D'autres facteurs peuvent influer de façon minime sur le champ de pesanteur local (Voir
Gravimétrie) :
• La composition du sous-sol (roches, grottes…)
L'accélération de la pesanteur peut se calculer comme suit :
g=9,780318 [m/s2] × (1 + 5,3024×10–3 × sin2(L) + 5,9×10–6 × sin2(2×L) – 3,15×10–7 × h)
où :
• L=la latitude
• h=l'altitude en mètre.
Au niveau de la mer, h=0 m :
• à l'équateur (L=0°) : g=9,7803 m/s²
• à la latitude (L=45°) : g=9,8063 m/s²
• aux pôles (L=90°) : g=9,8322 m/s²
La force de pesanteur englobe donc à la fois la gravité et la force centrifuge, qui elle provient du
mouvement de rotation. Il reste une force provenant de la rotation de la Terre qui ne peut être incluse dans la
pesanteur : la force de Coriolis, qui elle dépend de la vitesse de l'objet sur Terre. La pesanteur a été définie
pour être indépendante du mouvement de l'objet sur Terre.

Satellites de la Terre [modifier]

Article connexe : Lune.
Représentation à l'échelle de la taille et de la distance de la Terre et de la Lune. (1 px = 500 km)
La Lune est un satellite naturel, situé à environ 380 500 km de la Terre. Relativement grand, son
diamètre est environ le quart de celui de la Terre. Au sein du système solaire, c'est l'un des plus grands
satellites naturels (après Ganymède, Titan, Callisto et Io), le plus grand d'une planète non gazeuse. Elle est
relativement proche de la taille de la planète Mercure. Les satellites naturels orbitant autour des autres
planètes sont communément appelés « lunes » en référence à la Lune de la Terre.
L'attraction gravitationnelle entre la Terre et la Lune cause les marées sur Terre. Le même effet a
lieu sur la Lune, faisant en sorte que sa période de rotation est identique au temps qu'il lui faut pour orbiter
autour de la Terre, présentant ainsi toujours la même face vers notre planète. En orbitant autour de la Terre,
différentes parties du côté visible de la Lune sont illuminées par le Soleil, causant les phases lunaires.
À cause du couple des marées, la Lune s'éloigne de la Terre à un rythme d'environ 38 mm par an,
produisant aussi l'allongement du jour terrestre de 23 microsecondes par an. Sur plusieurs millions d'années,
l'effet cumulé de ces petites modifications produit d'importants changements. Durant la période du Dévonien,
à approximativement 410 millions d'années d'aujourd'hui, il y avait 400 jours dans une année, chaque jour
durant 21,8 heures.
Vue de la Terre, la Lune est assez éloignée pour avoir la même taille apparente que le Soleil. La
taille angulaire des deux corps est quasiment égale car même si le diamètre du Soleil est 400 fois plus grand
que celui de la Lune, celle-ci est 400 fois plus rapprochée de la Terre que ce dernier. Ceci permet encore
tout juste les éclipses totales sur Terre.
La théorie acceptée sur les origines de la Lune est celle d'un impact géant entre un planètoïde de la
taille de Mars, appelé Théia, et la Terre nouvellement formée. Cette hypothèse explique en partie le fait que
la composition de la Lune ressemble particulièrement à celle de la croûte terrestre.
La Terre a aussi deux satellites co-orbitaux, l'astéroïde (3753) Cruithne et 2002 AA29, ainsi que des
quasi-satellites, 2003 YN107 (jusqu'en 2006), 2001 GO2 et (164207) 2004 GU9.

Position dans l'Univers [modifier]

On sait aujourd'hui que la Terre tourne sur elle-même et autour du Soleil. Mais cette formulation
sous-entend un certain nombre de principes liés au développement de l'astronomie et de la physique, et qui
ont mis plusieurs siècles avant de s'affirmer.
La position de la Terre dans l'Univers fut la source de longs débats opposant durant des siècles
philosophes, savants et religieux de tous bords. Pendant longtemps, la Terre fut considérée comme au
centre de l'Univers, conception défendue par Aristote ou Ptolémée. Cette théorie, le géocentrisme, affirmait
que tous les objets célestes, Soleil, Lune, planètes et étoiles, (astres) orbitaient autour de la Terre.
L'héliocentrisme présente une Terre en orbite autour du Soleil, avec la Lune, satellite naturel de la Terre.
Défendue par Aristarque de Samos, cette théorie fut oubliée jusqu'à ce que Nicolas Copernic la redécouvre
et la complète dans son traité publié en 1543 : De revolutionibus orbium coelestium libri VI ). Tycho Brahe
proposa un système dans lequel le Soleil tournait autour de la Terre, et les autres planètes autour du Soleil.
L'idée de Copernic fut soutenue par Johannes Kepler et Galilée. Les ellipses képlériennes y firent beaucoup
en permettant des tables d'une précision jamais égalée, et en expliquant les variations de latitude des
planètes par rapport au plan de l'écliptique. À la fin du XVIIème, Isaac Newton introduisit sa mécanique qui
explique le mouvement de la Terre par les forces s'appliquant sur elle, en se plaçant dans un espace qu'il
supposait absolu. On disposait alors d'un système mécanique expliquant les mouvements des planètes,
conformes à la théorie héliocentrique.
Dans la modélisation actuelle de la mécanique newtonienne, on ne retient plus l'idée d'espace
absolu. La position des objets est définie par rapport à un référentiel, notion déjà présente chez Galilée,
Huygens et Newton, et la mécanique newtonienne repose sur l'hypothèse de l'existence d'une classe de
référentiels privilégiés, les référentiels galiléens. Dans un référentiel galiléen, les mouvements des planètes
s'expliquent par les seules forces de gravitation.
Dans le langage courant, quand rien n'est précisé, on se réfère à un référentiel galiléen. Cela est dû
au statut privilégié des référentiels galiléens en mécanique. C'est pour cela que la phrase « la Terre tourne »
n'est pas considérée comme imprécise en physique.
Il faut remarquer qu'en vertu du principe de relativité, y compris dans la version de Galilée, les effets
induits par le mouvement de la Terre sont extrêmement faibles pour ceux qui s'y trouvent. C'est pour cela
que les Anciens avaient pu supposer la Terre immobile.
Plus généralement les astrophysiciens considèrent aujourd'hui que non seulement l'Univers est
dépourvu de centre, mais même de tout point privilégié. Cette conjecture est appelée principe de Copernic,
puisque le nom de Copernic est associé à la fin de la vision de la Terre comme centre de l'univers.
Le drapeau non-officiel du Jour de la Terre.

Jour de la Terre [modifier]

Le Jour de la Terre a lieu le 22 avril, depuis 1970. Il marque la création de la mouvance écologiste.

Notes [modifier]
1. ↑ La Bille bleue, photo prise par l'équipage d'Apollo 17 le 7 décembre 1972. L'année 2009
marque le 50eanniversaire de la première photographie couleur à avoir été envoyée de l'espace le
1er décembre(1959 en science).
2. ↑ La Terre est la seule planète tournant autour du Soleil qui ne soit pas dotée d'un nom
universel pour toutes les langues issues de la mythologie grecque ou romaine. En français toutefois,
« Terre » s'apparente à Terra, déesse romaine de la terre, Gaïa en grec.
3. ↑ L'ozone est une des variante moléculaire de l'oxygène O 3.
4. ↑ La rotation de Vénus étant rétrograde, l’inclinaison de son axe est supérieure à 90°. On
pourrait dire que son axe est incliné de « -2,64° ».
5. ↑ Il utilise en effet comme base de son calcul la distance d'Alexandrie à Syène en stades, la
longueur du « stade » étant approximative à l'époque et la distance d'Alexandrie à Syène ne pouvant
alors s'apprécier que par une durée de marche de chameaux (Ian Stewart et Jack Cohen, The
Science of Discworld)
6. ↑ Augustin d'Hippone émet par exemple des doutes sur la possibilité que soient habitées les
antipodes, ce qui montre que le concept d'antipodes était bien admis.

Références [modifier]
1. ↑ Température relevée à El Azizia, Libye, le 13 septembre 1922 : (en) Global Measured
Extremes of Temperature and Precipitation [archive], National Climatic Data Center. Mis en ligne le
20 août 2008
2. ↑ Température relevée le 21 juillet 1983 à Vostok, Antarctique : (ru) Budretsky, A.B.,
« New absolute minimum of air temperature », dans Bulletin of the Soviet Antarctic Expedition ,
Gidrometeoizdat, Leningrad, no 105, 1984 [ texte intégral [archive] ]
3. ↑ pourcentage pondéral
4. ↑ (en) David R. Williams, « Mercury Fact Sheet [archive] », novembre 2007, NASA,
National Space Science Data Center.
5. ↑ (en) David R. Williams, « Venus Fact Sheet [archive] », avril 2005, NASA, National
Space Science Data Center.
6. ↑ (en) David R. Williams, « Earth Fact Sheet [archive] », avril 2007, NASA, National
Space Science Data Center.
7. ↑ (en) David R. Williams, « Mars Fact Sheet [archive] », novembre 2007, NASA, National
Space Science Data Center.
8. ↑ cf par exemple Vitruve, De architectura, Livre IX, ch. VII, trad. Coignard (1673) : « J'ai
parlé ici des constellations dont les figures ont été formées dans le Ciel par l'esprit divin qui est
auteur de la Nature, ainsi que le philosophe Démocrite les a désignées ; j'entends seulement celles
qui se lèvent et qui se couchent en notre horizon. Car tout de même que celles qui sont au
septentrion, et qui, faisant leur cours autour du pôle septentrional, ne se couchent point et ne
passent jamais sous la terre, ainsi il y en a d'autres sous la terre qui tournent aussi autour du pôle
méridional, demeurant toujours cachées sans se lever jamais sur la terre ; ce qui fait qu'on ne sait
point quelle est leur figure ; comme il se prouve par l'étoile appelée Canopus, que nous ne
connaissons que par le rapport des marchands qui ont voyagé dans les extrémités de l'Egypte, et
jusqu'au terres qui sont au bout du monde. »
9. ↑ Muhammad Hamidullah. L'Islam et son impulsion scientifique originelle, Tiers-Monde,
1982, vol. 23, n° 92, pp. 789: «... Enfin, l'imam Abou Hanifah (m. 767) savait que la terre était
sphérique ... »
10.↑ (en) IAU WG on NSFA – 10 août 2009 [archive] « Current Best Estimates »

Voir aussi [modifier]

Sur les autres projets Wikimédia :
• Terre sur Wikimedia Commons (ressources multimédia)
• Terre sur le Wiktionnaire (dictionnaire universel)
• Terre sur Wikibooks (livres pédagogiques)

Articles connexes [modifier]

• Conceptions actuelles
• Monde
• Liste des pays du monde
• Liste des pays du monde par continent
• Géographie
• Cartographie
• Projection cartographique
• Sciences de la Terre et de l'Univers
• Écologie
• Développement durable
• Pollution
• Géologie
• Géodésie
• Géophysique
• Hypothèse Gaïa
• Structure interne du globe terrestre
• Cycle biogéochimique
• Astronomie
• Planète
• Un point bleu pâle
• Conceptions anciennes
• Ératosthène
• Figure de la Terre dans l'Antiquité
• Figure de la Terre au Moyen Âge
• Figure de la Terre à la Renaissance
• Différents modèles
• le sphéroïde de Clairaut
• le modèle ellipsoïdal
• Le modèle incluant la notion de gravitation universelle

Liens externes [modifier]

• (fr) GéoManips, les mouvements de la Terre, site CNRS/sagascience
• (fr) Observer et interagir en 3D avec la Terre: Voir le jour et la nuit polaires
• (fr) Caractéristiques de la Terre sur le site de l'IMCCE
• (fr)
• (fr)
• (fr) Informations cartes et statistiques
• (fr) Cours de géologie en ligne
• (en) Moteur de recherche de coordonnées géographiques
• (en) Logiciel Google Earth
• (en) Vidéo comparant la taille de la Terre à d'autres astres
Système solaire

Étoile Soleil

Mercure • Vénus • Terre • Mars • Jupiter • Saturne • Uranus

• Neptune

Planètes naines Cérès • Pluton • Haumea • Makemake • Éris

Sedna • Quaoar • (225088) 2007 OR10 • Charon • (84522)

Objets massifs de la ceinture de Kuiper
2002 TC302 • Orcus • Varuna • 2007 UK126 • 2005 QU182

Mercurienne • Vénusienne • Terrestres : Lune • Martiennes •

Joviennes : Io · Europe · Ganymède · Callisto • Saturniennes : Titan
Lunes et lunes astéroïdales
• Uraniennes • Neptuniennes : Triton • Plutonniennes •
Haumeainnes • Érisienne

Anneaux Joviens• Saturniens • Uraniens • Neptuniens • Plutoniens •

de Ganymède • de Callisto • d'Europe • de Rhéa

Astéroïdes (liste) : Pallas · Junon · Vesta • Comètes :

Petits corps
1P/Halley · 2P/Encke • Damocloïde • Météoroïdes
Vulcanoïde • Ceinture d’astéroïdes • Centaure • Ceinture de
Principales zones Kuiper • Disque d’objets épars • Ceinture intermédiaire • Nuage de
Hills • Nuage d’Oort

Héliosphère • Héliopause • Héliogaine • Formation et

Autres évolution du système solaire • Milieu interplanétaire • Planètes
hypothétiques• C/1992 J1

Liste des objets du système solaire classés par : taille • masse • distance au Soleil

• Portail de la géodésie et de la géophysique

• Portail des sciences de la Terre et de l’Univers

• Portail de l’astronomie

• Portail de la géographie

• Portail de la biologie
Ce document provient de « ».

Catégories : Terre | Planète tellurique | [+]


From Wikipedia, the free encyclopedia

Jump to: navigation, search

"Primordial soup" redirects here. For the board game, see Primordial Soup (board game).
"Origin of life" redirects here. For views on the origins of life outside the natural sciences, see
Creation myth.
Pre-Cambrian stromatolites in the Siyeh Formation, Glacier National Park. In 2002, William Schopf of
UCLA published a paper in the scientific journal Nature arguing that geological formations such as this
possess 3.5 Ga (billion years old) fossilized cyanobacteria microbes. If true, they would be evidence of the
earliest known life on earth.
In natural science, abiogenesis (pronounced /ˌeɪbaɪ.ɵˈdʒɛnɨsɪs/, AY-bye-oh-JEN-ə-siss) or biopoesis
is the study of how life on Earth could have arisen from inanimate matter. It should not be confused with
evolution, which is the study of how groups of already living things change over time, or with cosmogony,
which covers how the universe might have arisen. Most amino acids, often called "the building blocks of life",
can form via natural chemical reactions unrelated to life, as demonstrated in the Miller–Urey experiment and
similar experiments, which involved simulating some of the conditions of the early Earth, in a scientific
laboratory.[1] In all living things, these amino acids are organized into proteins, and the construction of these
proteins is mediated by nucleic acids. Which of these organic molecules first arose and how they formed the
first life is the focus of abiogenesis.
In any theory of abiogenesis, two aspects of life have to be accounted for: replication, and
metabolism. The question of which came first gave rise to different types of theories. In the beginning,
metabolism-first theories (Oparin coacervate) were proposed, and only later thinking gave rise to modern,
replication-first approach.
In modern, still somewhat limited understanding, the first living things on Earth are thought to be
single cell prokaryotes (which lack a cell nucleus), perhaps evolved from protobionts (organic molecules
surrounded by a membrane-like structure).[2] The oldest ancient fossil microbe-like objects are dated to be
3.5 Ga (billion years old), approximately one billion years after the formation of the Earth itself.[3][4] By
2.4 Ga, the ratio of stable isotopes of carbon, iron and sulfur shows the action of living things on inorganic
minerals and sediments[5][6] and molecular biomarkers indicate photosynthesis, demonstrating that life on
Earth was widespread by this time.[7][8]
The sequence of chemical events that led to the first nucleic acids is not known. Several hypotheses
about early life have been proposed, most notably the iron-sulfur world theory (metabolism without genetics)
and the RNA world hypothesis (RNA life-forms).
• 1 Conceptual history
• 1.1 Spontaneous generation
• 1.2 Pasteur and Darwin
• 1.3 "Primordial soup" theory
• 2 Early conditions
• 3 Current models
• 3.1 Origin of organic molecules
• 3.1.1 "Soup" theory today: Miller's
experiment and subsequent work
• Regarding the reducing
• Regarding monomer
• Regarding monomer
• Regarding further
• 3.1.2 The deep sea vent theory
• 3.1.3 Fox's experiments
• 3.1.4 Eigen's hypothesis
[edit] Conceptual history
[edit] Spontaneous generation
Main article: Spontaneous generation
Until the early 19th century, people generally believed in the ongoing spontaneous generation of
certain forms of life from non-living matter. This was paired with heterogenesis, the belief that one form of life
derives from a different form (e.g. bees from flowers).[9] Classical notions of abiogenesis, now more
precisely known as spontaneous generation, held that certain complex, living organisms are generated by
decaying organic substances. According to Aristotle it was a readily observable truth that aphids arise from
the dew which falls on plants, fleas from putrid matter, mice from dirty hay, crocodiles from rotting logs at the
bottom of bodies of water, and so on.[10]
In the 17th century, such assumptions started to be questioned; for example, in 1646, Sir Thomas
Browne published his Pseudodoxia Epidemica (subtitled Enquiries into Very many Received Tenets, and
Commonly Presumed Truths), which was an attack on false beliefs and "vulgar errors." His conclusions were
not widely accepted. For example, his contemporary, Alexander Ross wrote: "To question this (i.e.,
spontaneous generation) is to question reason, sense and experience. If he doubts of this let him go to
Egypt, and there he will find the fields swarming with mice, begot of the mud of Nylus, to the great calamity of
the inhabitants."[11]
In 1665, Robert Hooke published the first drawings of a microorganism. Hooke was followed in 1676
by Anton van Leeuwenhoek, who drew and described microorganisms that are now thought to have been
protozoa and bacteria.[12] Many felt the existence of microorganisms was evidence in support of
spontaneous generation, since microorganisms seemed too simplistic for sexual reproduction, and asexual
reproduction through cell division had not yet been observed.
The first solid evidence against spontaneous generation came in 1668 from Francesco Redi, who
proved that no maggots appeared in meat when flies were prevented from laying eggs. It was gradually
shown that, at least in the case of all the higher and readily visible organisms, the previous sentiment
regarding spontaneous generation was false. The alternative seemed to be biogenesis: that every living thing
came from a pre-existing living thing (omne vivum ex ovo, Latin for "every living thing from an egg").
In 1768, Lazzaro Spallanzani demonstrated that microbes were present in the air, and could be killed
by boiling. In 1861, Louis Pasteur performed a series of experiments which demonstrated that organisms
such as bacteria and fungi do not spontaneously appear in sterile, nutrient-rich media.
[edit] Pasteur and Darwin

Charles Darwin in 1879.

By the middle of the 19th century, the theory of biogenesis had accumulated so much evidential
support, due to the work of Louis Pasteur and others, that the alternative theory of spontaneous generation
had been effectively disproven. Pasteur himself remarked, after a definitive finding in 1864, "Never will the
doctrine of spontaneous generation recover from the mortal blow struck by this simple experiment."[13] The
collapse of spontaneous generation, however, left a vacuum of scientific thought on the question of how life
had first arisen.
In a letter to Joseph Dalton Hooker on February 1, 1871,[14] Charles Darwin addressed the question,
suggesting that the original spark of life may have begun in a "warm little pond, with all sorts of ammonia and
phosphoric salts, lights, heat, electricity, etc. present, so that a protein compound was chemically formed
ready to undergo still more complex changes". He went on to explain that "at the present day such matter
would be instantly devoured or absorbed, which would not have been the case before living creatures were
formed."[15] In other words, the presence of life itself makes the search for the origin of life dependent on the
sterile conditions of the laboratory.
[edit] "Primordial soup" theory

Alexander Oparin (right) at the laboratory.

Further information: Miller–Urey experiment
No new notable research or theory on the subject appeared until 1924, when Alexander Oparin
reasoned that atmospheric oxygen prevents the synthesis of certain organic compounds that are necessary
building blocks for the evolution of life. In his The Origin of Life,[16][17] Oparin proposed that the
"spontaneous generation of life" that had been attacked by Louis Pasteur, did in fact occur once, but was
now impossible because the conditions found in the early earth had changed, and the presence of living
organisms would immediately consume any spontaneously generated organism. Oparin argued that a
"primeval soup" of organic molecules could be created in an oxygen-less atmosphere through the action of
sunlight. These would combine in ever-more complex fashions until they formed coacervate droplets. These
droplets would "grow" by fusion with other droplets, and "reproduce" through fission into daughter droplets,
and so have a primitive metabolism in which those factors which promote "cell integrity" survive, and those
that do not become extinct. Many modern theories of the origin of life still take Oparin's ideas as a starting
Around the same time, J. B. S. Haldane suggested that the Earth's pre-biotic oceans–very different
from their modern counterparts–would have formed a "hot dilute soup" in which organic compounds could
have formed. This idea was called biopoiesis or biopoesis, the process of living matter evolving from self-
replicating but nonliving molecules.[18][19]

[edit] Early conditions

Main article: Timeline of evolution
Morse and MacKenzie have suggested that oceans may have appeared first in the Hadean eon, as
soon as two hundred million years (200 Ma) after the Earth was formed, in a hot 100 °C (212 °F) reducing
environment, and that the pH of about 5.8 rose rapidly towards neutral.[20] This has been supported by
Wilde[3] who has pushed the date of the zircon crystals found in the metamorphosed quartzite of Mount
Narryer in Western Australia, previously thought to be 4.1–4.2 Ga, to 4.404 Ga. This means that oceans and
continental crust existed within 150 Ma of Earth's formation.
Despite this, the Hadean environment was one highly hazardous to life. Frequent collisions with large
objects, up to 500 kilometres (310 mi) in diameter, would have been sufficient to vaporise the ocean within a
few months of impact, with hot steam mixed with rock vapour leading to high altitude clouds completely
covering the planet. After a few months the height of these clouds would have begun to decrease but the
cloud base would still have been elevated for about the next thousand years. After that, it would have begun
to rain at low altitude. For another two thousand years rains would slowly have drawn down the height of the
clouds, returning the oceans to their original depth only 3,000 years after the impact event.[21]
Between 3.8 and 4.1 Ga, changes in the orbits of the gaseous giant planets may have caused a late
heavy bombardment that pockmarked the moon and other inner planets (Mercury, Mars, and presumably
Earth and Venus). This would likely have sterilized the planet had life appeared before that time.
By examining the time interval between such devastating environmental events, the time interval
when life might first have come into existence can be found for different early environments. The study by
Maher and Stevenson shows that if the deep marine hydrothermal setting provides a suitable site for the
origin of life, abiogenesis could have happened as early as 4.0 to 4.2 Ga, whereas if it occurred at the
surface of the earth abiogenesis could only have occurred between 3.7 and 4.0 Ga.[22]
Other research suggests a colder start to life. Work by Leslie Orgel and colleagues on the synthesis
of purines has shown that freezing temperatures are advantageous, due to the concentrating effect for key
precursors such as HCN.[23] Research by Stanley Miller and colleagues suggested that while adenine and
guanine require freezing conditions for synthesis, cytosine and uracil may require boiling temperatures.[24]
Based on this research, Miller suggested a beginning of life involving freezing conditions and exploding
meteorites.[25] An article in Discover Magazine points to research by the Miller group indicating the formation
of seven different amino acids and 11 types of nucleobases in ice when ammonia and cyanide were left in a
freezer from 1972–1997.[26][27] This article also describes research by Christof Biebricher showing the
formation of RNA molecules 400 bases long under freezing conditions using an RNA template, a single-
strand chain of RNA that guides the formation of a new strand of RNA. As that new RNA strand grows, it
adheres to the template.[28] The explanation given for the unusual speed of these reactions at such a low
temperature is eutectic freezing. As an ice crystal forms, it stays pure: only molecules of water join the
growing crystal, while impurities like salt or cyanide are excluded. These impurities become crowded in
microscopic pockets of liquid within the ice, and this crowding causes the molecules to collide more often.
Evidence of the early appearance of life comes from the Isua supercrustal belt in Western Greenland
and from similar formations in the nearby Akilia Islands. Carbon entering into rock formations has a ratio of
Carbon-13 (13C) to Carbon-12 (12C) of about −5.5 (in units of δ13C), where because of a preferential biotic
uptake of 12C, biomass has a δ13C of between −20 and −30. These isotopic fingerprints are preserved in the
sediments, and Mojzis has used this technique to suggest that life existed on the planet already by 3.85
billion years ago.[29] Lazcano and Miller (1994) suggest that the rapidity of the evolution of life is dictated by
the rate of recirculating water through mid-ocean submarine vents. Complete recirculation takes 10 million
years, thus any organic compounds produced by then would be altered or destroyed by temperatures
exceeding 300 °C (572 °F). They estimate that the development of a 100 kilobase genome of a DNA/protein
primitive heterotroph into a 7000 gene filamentous cyanobacterium would have required only 7 Ma.[30]

[edit] Current models

There is no truly "standard model" of the origin of life. Most currently accepted models draw at least
some elements from the framework laid out by the Oparin-Haldane hypothesis. Under that umbrella,
however, are a wide array of disparate discoveries and conjectures such as the following, listed in a rough
order of postulated emergence:
1. Some theorists suggest that the atmosphere of the early Earth may have been chemically
reducing in nature, composed primarily of methane (CH4), ammonia (NH3), water (H2O), hydrogen
sulfide (H2S), carbon dioxide (CO2) or carbon monoxide (CO), and phosphate (PO43-), with
molecular oxygen (O2) and ozone (O3) either rare or absent.
2. In such a reducing atmosphere, electrical activity can catalyze the creation of certain basic
small molecules (monomers) of life, such as amino acids. This was demonstrated in the Miller–Urey
experiment by Stanley L. Miller and Harold C. Urey in 1953.
3. Phospholipids (of an appropriate length) can spontaneously form lipid bilayers, a basic
component of the cell membrane.
4. A fundamental question is about the nature of the first self-replicating molecule. Since
replication is accomplished in modern cells through the cooperative action of proteins and nucleic
acids, the major schools of thought about how the process originated can be broadly classified as
"proteins first" and "nucleic acids first".
5. The principal thrust of the "nucleic acids first" argument is as follows:
1. The polymerization of nucleotides into random RNA molecules might have resulted
in self-replicating ribozymes (RNA world hypothesis)
2. Selection pressures for catalytic efficiency and diversity might have resulted in
ribozymes which catalyse peptidyl transfer (hence formation of small proteins), since
oligopeptides complex with RNA to form better catalysts. The first ribosome might have been
created by such a process, resulting in more prevalent protein synthesis.
3. Synthesized proteins might then outcompete ribozymes in catalytic ability, and
therefore become the dominant biopolymer, relegating nucleic acids to their modern use,
predominantly as a carrier of genomic information.
As of 2010, no one has yet synthesized a "protocell" using basic components which would have the
necessary properties of life (the so-called "bottom-up-approach"). Without such a proof-of-principle,
explanations have tended to be short on specifics. However, some researchers are working in this field,
notably Steen Rasmussen at Los Alamos National Laboratory and Jack Szostak at Harvard University.
Others have argued that a "top-down approach" is more feasible. One such approach, attempted by Craig
Venter and others at The Institute for Genomic Research, involves engineering existing prokaryotic cells with
progressively fewer genes, attempting to discern at which point the most minimal requirements for life were
reached. The biologist John Desmond Bernal coined the term Biopoesis for this process[citation needed],
and suggested that there were a number of clearly defined "stages" that could be recognised in explaining
the origin of life.
• Stage 1: The origin of biological monomers
• Stage 2: The origin of biological polymers
• Stage 3: The evolution from molecules to cell
Bernal suggested that evolution may have commenced early, some time between Stage 1 and 2[31].

[edit] Origin of organic molecules

There are two possible sources of organic molecules on the early Earth:
1. Terrestrial origins–organic synthesis driven by impact shocks or by other energy sources
(such as ultraviolet light or electrical discharges) (eg.Miller's experiments)
2. Extraterrestrial origins–delivery by objects (e.g. carbonaceous chondrites) or gravitational
attraction of organic molecules or primitive life-forms from space
Recently, estimates of these sources suggest that the heavy bombardment before 3.5 Ga within the
early atmosphere made available quantities of organics comparable to those produced by other energy

[edit] "Soup" theory today: Miller's experiment and subsequent work

Biochemist Robert Shapiro has summarized the "Primordial Soup" theory of Oparin and Haldane in
its "mature form" as follows:[33]
1. The early Earth had a chemically reducing atmosphere, as discussed above.
2. This atmosphere, exposed to energy in various forms, produced simple organic compounds
3. These compounds accumulated in a "soup", which may have been concentrated at various
locations (Shorelines, oceanic vents etc.).
4. By further transformation, more complex organic polymers— and ultimately life— developed in
the soup.

[edit] Regarding the reducing atmosphere

Whether the mixture of gases used in the Miller–Urey experiment truly reflects the atmospheric
content of early Earth is a controversial topic. Other less reducing gases produce a lower yield and variety. It
was once thought that appreciable amounts of molecular oxygen were present in the prebiotic
atmosphere[citation needed], which would have essentially prevented the formation of organic molecules;
however, the current scientific consensus is that such was not the case. (See Oxygen catastrophe).
[edit] Regarding monomer formation
Main article: Miller experiment
One of the most important pieces of experimental support for the "soup" theory came in 1953. A
graduate student, Stanley Miller, and his professor, Harold Urey, performed an experiment that demonstrated
how organic molecules could have spontaneously formed from inorganic precursors, under conditions like
those posited by the Oparin-Haldane Hypothesis. The now-famous "Miller–Urey experiment" used a highly
reduced mixture of gases–methane, ammonia and hydrogen–to form basic organic monomers, such as amino
acids.[34] This provided direct experimental support for the second point of the "soup" theory as described
above, and it is around the remaining two points of the theory that much of the debate now centers.
Apart from the Miller–Urey experiment, described above, the next most important step in research on
prebiotic organic synthesis was the demonstration by John Oró that the nucleic acid purine base, adenine,
was formed by heating aqueous ammonium cyanide solutions.[35] In support of abiogenesis in eutectic ice
(see above), more recent work demonstrated the formation of s-triazines (alternative nucleobases),
pyrimidines (including cytosine and uracil), and adenine from urea solutions subjected to freeze-thaw cycles
under a reductive atmosphere (with spark discharges as an energy source).[36]

[edit] Regarding monomer accumulation

The "soup" theory relies on the assumption proposed by Darwin (see above) that in an environment
with no pre-existing life, organic molecules may have accumulated and provided an environment for chemical

[edit] Regarding further transformation

The spontaneous formation of complex polymers from abiotically generated monomers under the
conditions posited by the "soup" theory is not at all a straightforward process. Besides the necessary basic
organic monomers, compounds that would have prohibited the formation of polymers were formed in high
concentration during the Miller–Urey and Oró experiments. The Miller experiment, for example, produces
many substances that would undergo cross-reactions with the amino acids or terminate the peptide chain.
More fundamentally, it can be argued that the most crucial challenge unanswered by this theory is
how the relatively simple organic building blocks polymerise and form more complex structures, interacting in
consistent ways to form a protocell. For example, in an aqueous environment hydrolysis of
oligomers/polymers into their constituent monomers would be favored over the condensation of individual
monomers into polymers.

[edit] The deep sea vent theory

The deep sea vent, or hydrothermal vent, theory for the origin of life on Earth posits that life may
have begun at submarine hydrothermal vents, where hydrogen-rich fluids emerge from below the sea floor
and interface with carbon dioxide-rich ocean water. Sustained chemical energy in such systems is derived
from redox reactions, in which electron donors, such as molecular hydrogen, react with electron acceptors,
such as carbon dioxide (see iron-sulfur world theory).

[edit] Fox's experiments

In the 1950s and 1960s, Sidney W. Fox studied the spontaneous formation of peptide structures
under conditions that might plausibly have existed early in Earth's history. He demonstrated that amino acids
could spontaneously form small peptides. These amino acids and small peptides could be encouraged to
form closed spherical membranes, called protenoid microspheres, which show many of the basic
characteristics of 'life'.[37]

[edit] Eigen's hypothesis

In the early 1970s the problem of the origin of life was approached by Manfred Eigen and Peter
Schuster of the Max Planck Institute for Biophysical Chemistry. They examined the transient stages between
the molecular chaos and a self-replicating hypercycle in a prebiotic soup.[38]
In a hypercycle, the information storing system (possibly RNA) produces an enzyme, which catalyzes
the formation of another information system, in sequence until the product of the last aids in the formation of
the first information system. Mathematically treated, hypercycles could create quasispecies, which through
natural selection entered into a form of Darwinian evolution. A boost to hypercycle theory was the discovery
that RNA, in certain circumstances, forms itself into ribozymes, capable of catalyzing their own chemical
reactions.[39] However, these reactions are limited to self-excisions (in which a longer RNA molecule
becomes shorter), and much rarer small additions that are incapable of coding for any useful protein. The
hypercycle theory is further degraded since the hypothetical RNA would require the existence of complex
biochemicals such as nucleotides which are not formed under the conditions proposed by the Miller–Urey

[edit] Hoffmann's contributions

Geoffrey W. Hoffmann, a student of Eigen, contributed to the concept of life involving both replication
and metabolism emerging from catalytic noise. His contributions included showing that an early sloppy
translation machinery can be stable against an error catastrophe of the type that had been envisaged as
problematical by Leslie Orgel ("Orgel's paradox")[40][41] and calculations regarding the occurrence of a set
of required catalytic activities together with the exclusion of catalytic activities that would be disruptive. This is
called the stochastic theory of the origin of life.[42]

[edit] Wächtershäuser's hypothesis

Main article: iron-sulfur world theory
Deep-sea black smoker
Another possible answer to this polymerization conundrum was provided in 1980s by the German
chemist Günter Wächtershäuser, in his iron-sulfur world theory. In this theory, he postulated the evolution of
(bio)chemical pathways as fundamentals of the evolution of life. Moreover, he presented a consistent system
of tracing today's biochemistry back to ancestral reactions that provide alternative pathways to the synthesis
of organic building blocks from simple gaseous compounds.
In contrast to the classical Miller experiments, which depend on external sources of energy (such as
simulated lightning or UV irradiation), "Wächtershäuser systems" come with a built-in source of energy,
sulfides of iron and other minerals (e.g. pyrite). The energy released from redox reactions of these metal
sulfides is not only available for the synthesis of organic molecules, but also for the formation of oligomers
and polymers. It is therefore hypothesized that such systems may be able to evolve into autocatalytic sets of
self-replicating, metabolically active entities that would predate the life forms known today.
The experiment produced a relatively small yield of dipeptides (0.4% to 12.4%) and a smaller yield of
tripeptides (0.10%) but the authors also noted that: "under these same conditions dipeptides hydrolysed

[edit] Radioactive beach hypothesis

Zachary Adam at the University of Washington, Seattle, claims that stronger tidal processes from a
much closer moon may have concentrated grains of uranium and other radioactive elements at the high
water mark on primordial beaches where they may have been responsible for generating life's building
blocks.[44] According to computer models reported in Astrobiology,[45] a deposit of such radioactive
materials could show the same self-sustaining nuclear reaction as that found in the Oklo uranium ore seam in
Gabon. Such radioactive beach sand provides sufficient energy to generate organic molecules, such as
amino acids and sugars from acetonitrile in water. Radioactive monazite also releases soluble phosphate
into regions between sand-grains, making it biologically "accessible". Thus amino acids, sugars and soluble
phosphates can all be simultaneously produced, according to Adam. Radioactive actinides, then in greater
concentrations, could have formed part of organo-metallic complexes. These complexes could have been
important early catalysts to living processes.
John Parnell of the University of Aberdeen suggests that such a process could provide part of the
"crucible of life" on any early wet rocky planet, so long as the planet is large enough to have generated a
system of plate tectonics which brings radioactive minerals to the surface. As the early Earth is believed to
have had many smaller "platelets" it would provide a suitable environment for such processes.[46]

[edit] Thermodynamic Origin of Life: Ultraviolet and Temperature-Assisted

Replication (UVTAR) Model
Karo Michaelian of the National Autonomous University of Mexico (UNAM) points out that any model
for the origin of life must take into account the fact that life is an irreversible thermodynamic process which
arises and persists to produce entropy. Entropy production is not incidental to the process of life, but rather
the fundamental reason for its existence. Present day life augments the entropy production of Earth by
catalysing the water cycle through evapotranspiration.[47] Michaelian argues that if the thermodynamic
function of life today is to produce entropy through coupling with the water cycle, then this probably was its
function at its very beginnings. It turns out that both RNA and DNA when in water solution are very strong
absorbers and extremely rapid dissipaters of ultraviolet light within the 200 nm - 300 nm wavelength range,
just that high energy part of the sun's spectrum that could have penetrated the dense prebiotic atmosphere.
Cnossen et al.[48] have shown that the amount of UV light reaching the Earth's surface in the Archean could
have been up to 31 orders of magnitude larger than it is today at 260 nm where RNA and DNA absorb most
strongly. Absorption and dissipation of UV light by these organic molecules at the Archean ocean surface
would have increased significantly the temperature of the surface skin layer leading to enhanced evaporation
and thus augmenting the primitive water cycle. Since absorption and dissipation of high energy photons is an
entropy producing process, Michaelian argues that non-equilbrium abiogenic synthesis of RNA and DNA
utilizing UV light [49] would have been thermodynamically favored.
A simple mechanism to explain the replication of RNA and DNA without the use of enzymes can also
be given within the same thermodynamic framework by assuming that life arose when the temperature of the
primitive seas had cooled to somewhat below the denaturing temperature of RNA or DNA (based on the ratio
of 18O/16O found in cherts of the Barberton greenstone belt of South Africa of about 3.5 to 3.2 Ga., surface
temperatures are predicted to have been around 70±15 °C [50], similar to RNA or DNA denaturing
temperatures). During the night, the surface water temperature would be below the denaturing temperature
and single strand RNA/DNA could act as a template for the formation of double strand RNA/DNA. During the
daylight hours, RNA and DNA would absorb UV light and convert this directly to heating of the ocean surface,
raising the local temperature enough to allow for denaturing of RNA and DNA. The copying process would be
repeated during the cool period overnight [51] . Such a temperature assisted mechanism of replication bears
similarity to Polymerase Chain Reaction (PCR), a routine laboratory procedure to multiply DNA segments.
Michaelian suggests that traditional origin of life research, expecting to describe the emergence of life from
near-equilibrium conditions, is erroneous and that non-equilibrium conditions must be considered, in
particular, the importance of entropy production to the emergence of life.
Since denaturation would be most probable in the late afternoon when the Archean sea surface
temperature would be highest, and since late afternoon sunlight is somewhat circularly polarized, the
homochirality of the organic molecules of life can also be explained within the proposed thermodynamic
[edit] Models to explain homochirality
Main article: Homochirality
Some process in chemical evolution must account for the origin of homochirality, i.e. all building
blocks in living organisms having the same "handedness" (amino acids being left-handed, nucleic acid
sugars (ribose and deoxyribose) being right-handed, and chiral phosphoglycerides). Chiral molecules can be
synthesized, but in the absence of a chiral source or a chiral catalyst, they are formed in a 50/50 mixture of
both enantiomers. This is called a racemic mixture. Clark has suggested that homochirality may have started
in space, as the studies of the amino acids on the Murchison meteorite showed L-alanine to be more than
twice as frequent as its D form, and L-glutamic acid was more than 3 times prevalent than its D counterpart. It
is suggested that polarised light has the power to destroy one enantiomer within the proto-planetary disk.
Noyes[53] showed that beta decay caused the breakdown of D-leucine, in a racemic mixture, and that the
presence of 14C, present in larger amounts in organic chemicals in the early Earth environment, could have
been the cause. Robert M. Hazen reports upon experiments conducted in which various chiral crystal
surfaces act as sites for possible concentration and assembly of chiral monomer units into macromolecules.
[54] Once established, chirality would be selected for.[55] Work with organic compounds found on meteorites
tends to suggest that chirality is a characteristic of abiogenic synthesis, as amino acids show a left-handed
bias, whereas sugars show a predominantly right-handed bias.[56]

[edit] Self-organization and replication

Main article: Self-organization
While features of self-organization and self-replication are often considered the hallmark of living
systems, there are many instances of abiotic molecules exhibiting such characteristics under proper
conditions. For example Martin and Russel show that physical compartmentation by cell membranes from the
environment and self-organization of self-contained redox reactions are the most conserved attributes of
living things, and they argue therefore that inorganic matter with such attributes would be life's most likely last
common ancestor.[57]
Virus self-assembly within host cells has implications for the study of the origin of life,[58] as it lends
further credence to the hypothesis that life could have started as self-assembling organic molecules.[59] [60]

[edit] From organic molecules to protocells

The question "How do simple organic molecules form a protocell?" is largely unanswered but there
are many hypotheses. Some of these postulate the early appearance of nucleic acids ("genes-first") whereas
others postulate the evolution of biochemical reactions and pathways first ("metabolism-first"). Recently,
trends are emerging to create hybrid models that combine aspects of both.

[edit] "Genes first" models: the RNA world

Main article: RNA world hypothesis
The RNA world hypothesis describes an early Earth with self-replicating and catalytic RNA but no
DNA or proteins. This has spurred scientists to try to determine if relatively short RNA molecules could have
spontaneously formed that were capable of catalyzing their own continuing replication.[61] A number of
hypotheses of modes of formation have been put forward.[62] Early cell membranes could have formed
spontaneously from proteinoids, protein-like molecules that are produced when amino acid solutions are
heated–when present at the correct concentration in aqueous solution, these form microspheres which are
observed to behave similarly to membrane-enclosed compartments. Other possibilities include systems of
chemical reactions taking place within clay substrates or on the surface of pyrite rocks. Factors supportive of
an important role for RNA in early life include its ability to act both to store information and catalyse chemical
reactions (as a ribozyme); its many important roles as an intermediate in the expression and maintenance of
the genetic information (in the form of DNA) in modern organisms; and the ease of chemical synthesis of at
least the components of the molecule under conditions approximating the early Earth. Relatively short RNA
molecules which can duplicate others have been artificially produced in the lab.[63] Such replicase RNA,
which functions as both code and catalyst provides a template upon which copying can occur. Jack Szostak
has shown that certain catalytic RNAs can, indeed, join smaller RNA sequences together, creating the
potential, in the right conditions for self-replication. If these were present, Darwinian selection would favour
the proliferation of such self-catalysing structures, to which further functionalities could be added.[64] Lincoln
and Joyce identified an RNA enzyme capable of self sustained replication.[65]
Researchers have pointed out difficulties for the abiotic synthesis of nucleotides from cytosine and
uracil.[66] Cytosine has a half-life of 19 days at 100 °C (212 °F) and 17,000 years in freezing water.[67]
Larralde et al., say that "the generally accepted prebiotic synthesis of ribose, the formose reaction, yields
numerous sugars without any selectivity."[68] and they conclude that their "results suggest that the backbone
of the first genetic material could not have contained ribose or other sugars because of their instability." The
ester linkage of ribose and phosphoric acid in RNA is known to be prone to hydrolysis.[69]
A slightly different version of the RNA-world hypothesis is that a different type of nucleic acid, such
as PNA, TNA or GNA, was the first one to emerge as a self-reproducing molecule, to be replaced by RNA
only later.[70][71] Pyrimidine ribonucleosides and their respective nucleotides have been prebiotically
synthesised by a sequence of reactions which by-pass the free sugars, and are assembled in a stepwise
fashion by going against the dogma that nitrogenous and oxygenous chemistries should be avoided. In a
series of publications, The Sutherland Group at the School of Chemistry, University of Manchester have
demonstrated high yielding routes to cytidine and uridine ribonucleotides built from small 2 and 3 carbon
fragments such as glycolaldehyde, glyceraldehyde or glyceraldehyde-3-phosphate, cyanamide and
cyanoacetylene. One of the steps in this sequence allows the isolation of enantiopure ribose aminooxazoline
if the enantiomeric excess of glyceraldehyde is 60 % or greater.[72] This can be viewed as a prebiotic
purification step, where the said compound spontaneously crystallised out from a mixture of the other
pentose aminooxazolines. Ribose aminooxazoline can then react with cyanoacetylene in a mild and highly
efficient manner to give the alpha cytidine ribonucleotide. Photoanomerization with UV light allows for
inversion about the 1' anomeric centre to give the correct beta stereochemistry.[73] In 2009 they showed that
the same simple building blocks allow access, via phosphate controlled nucleobase elaboration, to 2',3'-
cyclic pyrimidine nucleotides directly, which are known to be able to polymerise into RNA. This paper also
highlights the possibility for the photo-sanitization of the pyrimidine-2',3'-cyclic phosphates.[49] James
Ferris's studies have shown that clay minerals of montmorillonite will catalyze the formation of RNA in
aqueous solution, by joining activated mono RNA nucleotides to join together to form longer chains.[74]
Although these chains have random sequences, the possibility that one sequence began to non-randomly
increase its frequency by increasing the speed of its catalysis is possible to "kick start" biochemical evolution.

[edit] "Metabolism first" models

Several models reject the idea of the self-replication of a "naked-gene" and postulate the emergence
of a primitive metabolism which could provide an environment for the later emergence of RNA replication.

[edit] Iron-sulfur world

One of the earliest incarnations of this idea was put forward in 1924 with Alexander Oparin's notion of
primitive self-replicating vesicles which predated the discovery of the structure of DNA. More recent variants
in the 1980s and 1990s include Günter Wächtershäuser's iron-sulfur world theory and models introduced by
Christian de Duve based on the chemistry of thioesters. More abstract and theoretical arguments for the
plausibility of the emergence of metabolism without the presence of genes include a mathematical model
introduced by Freeman Dyson in the early 1980s and Stuart Kauffman's notion of collectively autocatalytic
sets, discussed later in that decade.
However, the idea that a closed metabolic cycle, such as the reductive citric acid cycle, could form
spontaneously (proposed by Günter Wächtershäuser) remains debated. In an article entitled "Self-Organizing
Biochemical Cycles",[75] the late Leslie Orgel summarized his analysis of the proposal by stating, "There is
at present no reason to expect that multistep cycles such as the reductive citric acid cycle will self-organize
on the surface of FeS/FeS2 or some other mineral." It is possible that another type of metabolic pathway was
used at the beginning of life. For example, instead of the reductive citric acid cycle, the "open" acetyl-CoA
pathway (another one of the five recognised ways of carbon dioxide fixation in nature today) would be
compatible with the idea of self-organisation on a metal sulfide surface. The key enzyme of this pathway,
carbon monoxide dehydrogenase/acetyl-CoA synthase harbours mixed nickel-iron-sulfur clusters in its
reaction centers and catalyses the formation of acetyl-CoA (which may be regarded as a modern form of
acetyl-thiol) in a single step.

[edit] Thermosynthesis world

Today’s bioenergetic process of fermentation is related to the just mentioned citric acid cycle or the
Acetyl-CoA pathway that have been connected to the primordial iron-sulfur world. In a different approach,
today’s bioenergetic process of chemiosmosis, which plays an essential role in cellular respiration and
photosynthesis, is considered as more fundamental than fermentation: in Anthonie Muller’s “thermosynthesis
world” the ATP Synthase enzyme that sustains chemiosmosis is proposed as today’s enzyme that is the
closest connected to the first metabolic process.[76][77]
First life needed an energy source to bring about the condensation reaction that yielded the peptide
bonds of proteins and the phosphodiester bonds of RNA. In a generalization and thermal variation of the
binding change mechanism of today’s ATP Synthase, the “First Protein” would have bound substrates
(peptides, phosphate, nucleosides, RNA ‘monomers’) and condensed them to a reaction product that
remained bound until it after a temperature change was released upon a thermal unfolding.
The energy source of the thermosynthesis world was thermal cycling, the result of suspension of the
protocell in a convection current, as is plausible in a volcanic hot spring; the convection accounts for the self-
organization and dissipative structure required in any origin of life model. The still ubiquitous role of thermal
cycling in germination and cell division is considered a relic of primordial thermosynthesis.
By phosphorylating cell membrane lipids, this ‘First Protein’ gave a selective advantage to the lipid
protocell that contained the protein. In the beginning this First Protein also synthesized a library with many
proteins, of which only a minute fraction had thermosynthesis capabilities. Just as proposed by Dyson [78] for
the first proteins, the First Protein propagated functionally: it made daughters with similar capabilities, but it
did not copy itself. Functioning daughters consisted of different amino acid sequences.
Over a long time, RNA sequences were selected among the at first randomly synthesized RNAs by
the criterion of speed and efficiency increase of First Protein synthesis, for instance by the creation of RNA
that functioned as messenger RNA[79], Transfer RNA[80] and ribosomal RNA, or, even more generally, all
the components of the RNA World were also generated and selected. The thermosynthesis world therefore in
theory accounts for the origin of the genetic machinery.
Whereas the iron-sulfur world identifies a circular pathway as the most simple—and therefore
assumes the existence of enzymes—the thermosynthesis world does not even invoke a pathway, and does
not assume the existence of regular enzymes: ATP Synthase’s binding change mechanism resembles a
physical adsorption process that yields free energy[81], rather than a regular enzyme’s mechanism, which
decreases the free energy. The RNA World also implies the existence of several enzymes. But even the
emergence of a single enzyme by chance is implausible.[82] The thermosynthesis world is therefore more
simple, and thus more plausible, than the iron-sulfur and RNA worlds.

[edit] Possible role of bubbles

Waves breaking on the shore create a delicate foam composed of bubbles. Winds sweeping across
the ocean have a tendency to drive things to shore, much like driftwood collecting on the beach. It is possible
that organic molecules were concentrated on the shorelines in much the same way. Shallow coastal waters
also tend to be warmer, further concentrating the molecules through evaporation. While bubbles composed
mostly of water burst quickly, water containing amphiphiles forms much more stable bubbles, lending more
time to the particular bubble to perform these crucial reactions.
Amphiphiles are oily compounds containing a hydrophilic head on one or both ends of a hydrophobic
molecule. Some amphiphiles have the tendency to spontaneously form membranes in water. A spherically
closed membrane contains water and is a hypothetical precursor to the modern cell membrane. If a protein
would increase the integrity of its parent bubble, that bubble had an advantage, and was placed at the top of
the natural selection waiting list. Primitive reproduction can be envisioned when the bubbles burst, releasing
the results of the 'experiment' into the surrounding medium. Once enough of the 'right stuff' was released into
the medium, the development of the first prokaryotes, eukaryotes, and multicellular organisms could be
Similarly, bubbles formed entirely out of protein-like molecules, called microspheres, will form
spontaneously under the right conditions. But they are not a likely precursor to the modern cell membrane, as
cell membranes are composed primarily of lipid compounds rather than amino-acid compounds (for types of
membrane spheres associated with abiogenesis, see protobionts, micelle, coacervate).
A recent model by Fernando and Rowe[84] suggests that the enclosure of an autocatalytic non-
enzymatic metabolism within protocells may have been one way of avoiding the side-reaction problem that is
typical of metabolism first models.

[edit] Other models

[edit] Autocatalysis
In 1993 Stuart Kauffman proposed that life initially arose as autocatalytic chemical networks.[85]
British ethologist Richard Dawkins wrote about autocatalysis as a potential explanation for the origin
of life in his 2004 book The Ancestor's Tale. Autocatalysts are substances which catalyze the production of
themselves, and therefore have the property of being a simple molecular replicator. In his book, Dawkins
cites experiments performed by Julius Rebek and his colleagues at the Scripps Research Institute in
California in which they combined amino adenosine and pentafluorophenyl ester with the autocatalyst amino
adenosine triacid ester (AATE). One system from the experiment contained variants of AATE which
catalysed the synthesis of themselves. This experiment demonstrated the possibility that autocatalysts could
exhibit competition within a population of entities with heredity, which could be interpreted as a rudimentary
form of natural selection.
[edit] Clay theory
A model for the origin of life based on clay was forwarded by A. Graham Cairns-Smith of the
University of Glasgow in 1985 and explored as a plausible illustration by several other scientists, including
Richard Dawkins[86]. Clay theory postulates that complex organic molecules arose gradually on a pre-
existing, non-organic replication platform—silicate crystals in solution. Complexity in companion molecules
developed as a function of selection pressures on types of clay crystal is then exapted to serve the replication
of organic molecules independently of their silicate "launch stage".
Cairns-Smith is a staunch critic of other models of chemical evolution.[87] However, he admits that
like many models of the origin of life, his own also has its shortcomings (Horgan 1991).
In 2007, Kahr and colleagues reported their experiments to examine the idea that crystals can act as
a source of transferable information, using crystals of potassium hydrogen phthalate. "Mother" crystals with
imperfections were cleaved and used as seeds to grow "daughter" crystals from solution. They then
examined the distribution of imperfections in the crystal system and found that the imperfections in the
mother crystals were indeed reproduced in the daughters. The daughter crystals had many additional
imperfections. For a gene-like behavior the additional imperfections should be much less than the parent
ones, thus Kahr concludes that the crystals "were not faithful enough to store and transfer information from
one generation to the next".[88][89]

[edit] Gold's "Deep-hot biosphere" model

In the 1970s, Thomas Gold proposed the theory that life first developed not on the surface of the
Earth, but several kilometers below the surface. The discovery in the late 1990s of nanobes (filamental
structures that are smaller than bacteria, but that may contain DNA) in deep rocks [90] might be seen as
lending support to Gold's theory.
It is now reasonably well established that microbial life is plentiful at shallow depths in the Earth, up
to 5 kilometres (3.1 mi) below the surface,[90] in the form of extremophile archaea, rather than the better-
known eubacteria (which live in more accessible conditions). It is claimed that discovery of microbial life
below the surface of another body in our solar system would lend significant credence to this theory. Thomas
Gold also asserted that a trickle of food from a deep, unreachable, source is needed for survival because life
arising in a puddle of organic material is likely to consume all of its food and become extinct. Gold's theory is
that flow of food is due to out-gassing of primordial methane from the Earth's mantle; more conventional
explanations of the food supply of deep microbes (away from sedimentary carbon compounds) is that the
organisms subsist on hydrogen released by an interaction between water and (reduced) iron compounds in

[edit] "Primitive" extraterrestrial life

An alternative to Earthly abiogenesis is the hypothesis that primitive life may have originally formed
extraterrestrially, either in space or on a nearby planet (Mars). (Note that exogenesis is related to, but not the
same as, the notion of panspermia). A supporter of this theory was Francis Crick.
Organic compounds are relatively common in space, especially in the outer solar system where
volatiles are not evaporated by solar heating.[91] Comets are encrusted by outer layers of dark material,
thought to be a tar-like substance composed of complex organic material formed from simple carbon
compounds after reactions initiated mostly by irradiation by ultraviolet light. It is supposed that a rain of
material from comets could have brought significant quantities of such complex organic molecules to Earth.
An alternative but related hypothesis, proposed to explain the presence of life on Earth so soon after
the planet had cooled down, with apparently very little time for prebiotic evolution, is that life formed first on
early Mars. Due to its smaller size Mars cooled before Earth (a difference of hundreds of millions of years),
allowing prebiotic processes there while Earth was still too hot. Life was then transported to the cooled Earth
when crustal material was blasted off Mars by asteroid and comet impacts. Mars continued to cool faster and
eventually became hostile to the continued evolution or even existence of life (it lost its atmosphere due to
low volcanism); Earth is following the same fate as Mars, but at a slower rate.
Neither hypothesis actually answers the question of how life first originated, but merely shifts it to
another planet or a comet. However, the advantage of an extraterrestrial origin of primitive life is that life is
not required to have evolved on each planet it occurs on, but rather in a single location, and then spread
about the galaxy to other star systems via cometary and/or meteorite impact. Evidence to support the
plausibility of the concept is scant, but it finds support in recent study of Martian meteorites found in
Antarctica and in studies of extremophile microbes.[92] Additional support comes from a recent discovery of
a bacterial ecosytem whose energy source is radioactivity.[93]
A 2001 experiment led by Jason Dworkin[94] subjected a frozen mixture of water, methanol,
ammonia and carbon monoxide to UV radiation, mimicking conditions found in an extraterrestrial
environment. This combination yielded large amounts of organic material that self-organised to form bubbles
or micelles when immersed in water. Dworkin considered these bubbles to resemble cell membranes that
enclose and concentrate the chemistry of life, separating their interior from the outside world.
The bubbles produced in these experiments were between 10 to 40 micrometres (0.00039 to 0.0016
in), or about the size of red blood cells. Remarkably, the bubbles fluoresced, or glowed, when exposed to UV
light. Absorbing UV and converting it into visible light in this way was considered one possible way of
providing energy to a primitive cell. If such bubbles played a role in the origin of life, the fluorescence could
have been a precursor to primitive photosynthesis. Such fluorescence also provides the benefit of acting as a
sunscreen, diffusing any damage that otherwise would be inflicted by UV radiation. Such a protective function
would have been vital for life on the early Earth, since the ozone layer, which blocks out the sun's most
destructive UV rays, did not form until after photosynthetic life began to produce oxygen.[56]

[edit] Extraterrestrial amino acids

Another idea is that amino acids which were formed extra-terrestrially arrived on Earth via comets. In
2009 it was announced by NASA that scientists have identified one of the fundamental chemical buildings
blocks of life in a comet for the first time: glycine, an amino acid, was detected in the material ejected from
Comet Wild-2 in 2004 and grabbed by NASA's Stardust probe. Tiny grains, just a few thousandths of a
millimetre in size, were collected from the comet and returned to Earth in 2006 in a sealed capsule, and
distributed among the world's leading astro-biology labs. NASA said in a statement that it took some time for
the investigating team, led by Dr Jamie Elsila, to convince itself that the glycine signature found in Stardust's
sample bay was genuine and not just Earthly contamination. Glycine has been detected in meteorites before
and there are also observations in interstellar gas clouds claimed for telescopes, but the Stardust find is
described as a first in cometary material. It is known that prior to the emergence of life on Earth, the early
solar system's planets were regularly bombarded by comets. Dr. Carl Pilcher, who leads NASA's
Astrobiology Institute commented that "The discovery of glycine in a comet supports the idea that the
fundamental building blocks of life are prevalent in space, and strengthens the argument that life in the
Universe may be common rather than rare."[95]
[edit] Lipid World
This theory postulates that the first self-replicating object was lipid-like.[96] It is known that
phospholipids form bilayers in water while under agitation– the same structure as in cell membranes. These
molecules were not present on early Earth, however other amphiphilic long chain molecules also form
membranes. Furthermore, these bodies may expand (by insertion of additional lipids), and under excessive
expansion may undergo spontaneous splitting which preserves the same size and composition of lipids in the
two progenies. The main idea in this theory is that the molecular composition of the lipid bodies is the
preliminary way for information storage, and evolution led to the appearance of polymer entities such as RNA
or DNA that may store information favorably. Still, no biochemical mechanism has been offered to support
the Lipid World theory.

[edit] Polyphosphates
The problem with most scenarios of abiogenesis is that the thermodynamic equilibrium of amino acid
versus peptides is in the direction of separate amino acids. What has been missing is some force that drives
polymerization. The resolution of this problem may well be in the properties of polyphosphates.[97][98]
Polyphosphates are formed by polymerization of ordinary monophosphate ions PO 4−3. Several mechanisms
for such polymerization have been suggested. Polyphosphates cause polymerization of amino acids into
peptides[citation needed]. They are also logical precursors in the synthesis of such key biochemical
compounds as ATP. A key issue seems to be that calcium reacts with soluble phosphate to form insoluble
calcium phosphate (apatite), so some plausible mechanism must be found to keep calcium ions from causing
precipitation of phosphate. There has been much work on this topic over the years, but an interesting new
idea is that meteorites may have introduced reactive phosphorus species on the early Earth.[99]
[edit] PAH world hypothesis
Main article: PAH world hypothesis
Other sources of complex molecules have been postulated, including extraterrestrial stellar or
interstellar origin. For example, from spectral analyses, organic molecules are known to be present in comets
and meteorites. In 2004, a team detected traces of polycyclic aromatic hydrocarbons (PAH's) in a nebula.
[100] Those are the most complex molecules so far found in space. The use of PAH's has also been
proposed as a precursor to the RNA world in the PAH world hypothesis.[101] The Spitzer Space Telescope
has recently detected a star, HH 46-IR, which is forming by a process similar to that by which the sun formed.
In the disk of material surrounding the star, there is a very large range of molecules, including cyanide
compounds, hydrocarbons, and carbon monoxide. PAHs have also been found all over the surface of galaxy
M81, which is 12 million light years away from the Earth, confirming their widespread distribution in space.

[edit] Multiple genesis

Different forms of life may have appeared quasi-simultaneously in the early history of Earth.[103] The
other forms may be extinct, leaving distinctive fossils through their different biochemistry (e.g., using arsenic
instead of phosphorus), survive as extremophiles, or simply be unnoticed through their being analogous to
organisms of the current life tree. Hartman[104] for example combines a number of theories together, by
proposing that:
The first organisms were self-replicating iron-rich clays which fixed carbon dioxide into oxalic
and other dicarboxylic acids. This system of replicating clays and their metabolic phenotype
then evolved into the sulfide rich region of the hotspring acquiring the ability to fix nitrogen.
Finally phosphate was incorporated into the evolving system which allowed the synthesis of
nucleotides and phospholipids. If biosynthesis recapitulates biopoesis, then the synthesis of
amino acids preceded the synthesis of the purine and pyrimidine bases. Furthermore the
polymerization of the amino acid thioesters into polypeptides preceded the directed
polymerization of amino acid esters by polynucleotides.

Lynn Margulis's endosymbiotic theory suggests that multiple forms of bacteria entered into symbiotic
relationship to form the eukaryotic cell. The horizontal transfer of genetic material between bacteria promotes
such symbiotic relationships, and thus many separate organisms may have contributed to building what has
been recognised as the Last Universal Common Ancestor (LUCA) of modern organisms. James Lovelock's
Gaia theory, proposes that such bacterial symbiosis establishes the environment as a system produced by
and supportive of life. His arguments strongly weaken the case for life having evolved elsewhere in the solar

[edit] See also

• Astrochemistry
• Autocatalytic reactions and order creation
• Biogenesis
• Common descent
• Drake equation
• Entropy and life
• History of Earth
• List of independent discoveries
• List of publications in biology
• Mediocrity principle
• Origin of the world's oceans
• Mimivirus
• Planetary habitability
• Rare Earth hypothesis
• Shadow biosphere
• Thermosynthesis
• Zeolite
• John Needham

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[edit] Further reading
• Arrhenius, Gustaf; et al. (1997). "Entropy and Charge in Molecular Evolution—the Case of
Phosphate". Journal of Theoretical Biology 187 (4): 503–522. doi:10.1006/jtbi.1996.0385.
PMID 9299295.
• Buehler, Lukas K. (2000–2005) The physico-chemical basis of life, accessed 27 October 2005.
• Davies, Paul (1998). The Fifth Miracle. Penguin Science, London. ISBN 0-140-28226-2.
• De Duve, Christian (January 1996). Vital Dust: The Origin and Evolution of Life on Earth .
Basic Books. ISBN 0-465-09045-1.
• Fernando CT, Rowe, J (2007). "Natural selection in chemical evolution". Journal of
Theoretical Biology 247 (1): 152–67. doi:10.1016/j.jtbi.2007.01.028. PMID 17399743.
• Hartman, Hyman (1998). "Photosynthesis and the Origin of Life". Origins of Life and
Evolution of Biospheres 28 (4–6): 515–521. doi:10.1023/A:1006548904157.
• Harris, Henry (2002). Things come to life. Spontaneous generation revisited . Oxford: Oxford
University Press. ISBN 0198515383.
• Hazen, Robert M. (December 2005). Genesis: The Scientific Quest for Life's Origins. Joseph
Henry Press. ISBN 0-309-09432-1.
• Gribbon, John (1998). The Case of the Missing Neutrino's and other Curious Phenomena of
the Universe. Penguin Science, London. ISBN 0-140-28734-5.
• Horgan, J (1991). "In the beginning". Scientific American 264: 100–109. (Cited on p. 108).
• Huber, C. and Wächterhäuser, G., (1998). "Peptides by activation of amino acids with CO on
(Ni,Fe)S surfaces: implications for the origin of life". Science 281 (5377): 670–672.
doi:10.1126/science.281.5377.670. PMID 9685253. (Cited on p. 108).
• Knoll, Andrew H. (2003). Life on a Young Planet: The First Three Billion Years of Evolution
on Earth. Princeton University Press. ISBN 0691009783.
• Luisi, Pier Luigi (2006). The Emergence of Life: From Chemical Origins to Synthetic Biology .
Cambridge University Press. ISBN 0521821177.
• Martin, W. and Russell M.J. (2002). "On the origins of cells: a hypothesis for the evolutionary
transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to
nucleated cells". Philosophical Transactions of the Royal Society: Biological sciences 358 (1429): 59–
85. doi:10.1098/rstb.2002.1183. PMID 12594918.
• Maynard Smith, John; Szathmary, Eors (2000-03-16). The Origins of Life: From the Birth of
Life to the Origin of Language. Oxford Paperbacks. ISBN 0-19-286209-X.
• Morowitz, Harold J. (1992) "Beginnings of Cellular Life: Metabolism Recapitulates
Biogenesis". Yale University Press. ISBN 0-300-05483-1
• NASA Astrobiology Institute: Earth's Early Environment and Life
• NASA Specialized Center of Research and Training in Exobiology: Gustaf O. Arrhenius
• Pitsch, Stefan; Krishnamurthy, Ramanarayanan; Arrhenius, Gustaf (2000). "Concentration of
Simple Aldehydes by Sulfite-Containing Double-Layer Hydroxide Minerals: Implications for
Biopoesis" (abstract). Helvetica Chimica Acta 83 (9): 2398 2411. doi:10.1002/1522-
• Russell MJ, Hall AJ, Cairns-Smith AG, Braterman PS (1988). "Submarine hot springs and the
origin of life". Nature 336: 117. doi:10.1038/336117a0.
• Dedicated issue of Philosophical Transactions B on Major Steps in Cell Evolution freely
• Dedicated issue of Philosophical Transactions B on the Emergence of Life on the Early Earth
freely available.

[edit] External links

• The Deep Hot Biosphere Theory - Thomas Gold, article from PNAS Proc. Natl. Acad. Sci.
USA Vol. 89, pp. 6045-6049, July 1992 Microbiology
• The Origin of Life Video by John Maynard Smith
• "Harvard Team Creates the World's 1st Synthesized Cells"
• Martin M Hanczyc and Jack W Szostak. Replicating vesicles as models of primitive cell
growth and division. Current Opinion in Chemical Biology 2004, 8:660–664.PDF (192 KB)
• Martin A. Nowak and Hisashi Ohtsuki.Prevolutionary dynamics and the origin of evolution.
Proceedings of the National Academy of Sciences 2008
• "Exploring Life's Origins: a Virtual Exhibit"
• "SELF-REPLICATION: Even peptides do it" by Stuart A. Kauffman (web archive version as
original page no longer accessible)
• Origins of Life website including papers, resources, by Dr. Michael Russell at the U. of
• Possible Connections Between Interstellar Chemistry and the Origin of Life on the Earth
• Scientists Find Clues That Life Began in Deep Space—NASA Astrobiology Institute
• Self-organizing biochemical cycles—by Leslie Orgel
• How Life Began: New Research Suggests Simple Approach
• Primordial Soup's On: Scientists Repeat Evolution's Most Famous Experiment –an article in
Scientific American. March 28, 2007
• Illustrations from Evolution (textbook)
• An abiogenesis primer for laymen

Origin of life

Hypercycle · Protobiont · Universal common descent · Last universal ancestor · RNA world
hypothesis · Iron-sulfur world theory · PAH world hypothesis · Miller–Urey experiment · Panspermia

Retrieved from ""

Categories: Evolutionarily significant biological phenomena | Evolutionary biology | Origin of life


Origine de la vie
Un article de Wikipédia, l'encyclopédie libre.
(Redirigé depuis Origines de la vie)
Aller à : Navigation, rechercher
Cet article est consacré aux origines de la vie d'un point de vue scientifique. Les aspects mythiques
et religieux sont traités dans l'article Cosmogonie. La précédente théorie scientifique de l'origine de la vie est
traitée dans l'article génération spontanée.
Les origines de la vie sur Terre demeurent incertaines. Cependant, de nombreuses théories
scientifiques existent pour expliquer l'apparition de la vie, telle que nous la connaissons aujourd'hui, dont on
pense qu'elle remonte à environ 3,5 à 3,8 milliards d'années.
Cet article traite des événements antérieurs à l'apparition des trois grandes lignées du vivant.
Stromatolites du précambrien, dans la formation de Siyeh. En 2002, William Schopf a affirmé que
ces formations étaient âgées de 3,5 milliards d'années, elles seraient alors les plus anciennes traces de vie
sur Terre.
Stromatolites dans la Baie Shark (Australie).
• 1 L'apparition de la vie
• 1.1 L'origine des molécules organiques
• 1.1.1 Une explication : la condensation sur
surfaces minérales
• 1.1.2 L'expérience Urey-Miller et l'origine
des molécules organiques
• L'exploitation de l'idée de
• 1.1.3 L'expérience de Powner-Sutherland et
l'origine des nucléotides à base pyrimidique
• 1.1.4 Asymétrie des biomolécules
• Théories sur l'origine de
• Cristaux et énantiomères
• Asymétricité et
• Formation d'énantiomères
dans l'espace
• 1.2 Des molécules organiques aux protocellules
• 1.2.1 Compartiments isolés
L'apparition de la vie [modifier]
John Maynard Smith et Eörs Szathmáry définissent huit transitions majeures de l'évolution, dont trois
concernent l'apparition de la vie :
1. transition depuis des molécules autoréplicantes vers une population de molécules dans un
compartiment isolé ;
2. passage de réplicateurs indépendants aux chromosomes ;
3. transition d'un monde à ARN — où ce dernier joue le rôle de génome et d'enzyme — à un
monde à ADN et protéines.
John Desmond Bernal, quant à lui, suggère les trois étapes suivantes :
1. apparition de monomères organiques,
2. transitions vers des polymères organiques,
3. évolution depuis des molécules vers la cellule.
De fait, il n'existe pas de modèle « standard » pour décrire l'origine de la vie. Cependant le modèle le
plus couramment accepté est fondé sur l'enchaînement supposé des événements suivants :
1. Des conditions prébiotiques plausibles entraînent la création de molécules organiques
simples qui sont les briques de base du vivant.
2. Des phospholipides forment spontanément des doubles couches qui sont la structure de
base des membranes cellulaires.
3. Les mécanismes qui produisent aléatoirement des molécules d'ARN (acide ribonucléique),
en mesure d'agir comme des ARN-enzymes capables, dans certaines conditions très particulières,
de se dupliquer. C'est une première forme de génome, et nous sommes alors en présence de
4. Les ARN-enzymes sont progressivement remplacées par des protéines-enzymes, grâce à
l'apparition des ribozymes, ceux-ci étant capables de réaliser la synthèse des protéines.
5. L'ADN apparaît et remplace l'ARN dans le rôle de support du génome, dans le même temps
les ribozymes sont complétés par des protéines, formant les ribosomes. C'est l'apparition de
l'organisation actuelle des organismes vivants.
Les étapes 2 et 3 sont parfois inversées, l'isolement en compartiment étant alors présenté après
l'apparition des ARN autoréplicants.
L'origine des molécules organiques [modifier]

Une explication : la condensation sur surfaces minérales [modifier]

Cristaux de pyrite
L'assemblage de petites molécules (comme les acides aminés) en macromolécules (comme les
protéines) nécessite l'élimination de molécules d'eau. Or, la thermodynamique indique qu'il est défavorable
de réaliser une telle condensation dans l'eau elle-même. Il est possible pour résoudre cette contradiction de
faire appel à des surfaces minérales, comme les micas, les argiles ou les pyrites. L'adsorption des petites
molécules sur ces surfaces les concentre et les modifie chimiquement, ce qui peut rendre la formation de
macromolécules plus favorable.
L'argile, par exemple, se trouve très abondamment sur Terre et est constituée d'un empilement de
couches fines. Entre les différentes couches de l'argile peuvent se glisser certaines petites molécules
organiques, ce qui permet une adsorption importante. L'argile est aussi un catalyseur très efficace pour de
nombreuses réactions organiques, et aurait donc pu permettre la polymérisation des acides aminés et/ou
des acides nucléiques. Le chimiste anglais Cairns-Smith a développé cette hypothèse dans Seven clues to
the origin of life en 1985 (traduction française : L'énigme de la vie, 1990).

L'expérience Urey-Miller et l'origine des molécules organiques [modifier]

Article détaillé : Expérience de Miller-Urey.
En 1953, Stanley Miller, accompagné de Harold Urey, a voulu reproduire les conditions de la Terre
primitive. Ils ont enfermé dans un ballon des gaz (méthane CH 4, ammoniac NH3, hydrogène H2 et eau H2O)
et soumis le mélange à des décharges électriques pendant sept jours.
Ils ont obtenu des molécules organiques, les briques du vivant, et notamment de l'urée (CON2H4), du
formaldéhyde (H2CO), de l'acide cyanhydrique (HCN), des bases et des acides aminés (AA), certains
composés étant présents à plus de 2 %.
Miller et Urey ont utilisé une atmosphère réductrice, méthane CH4, NH3, H2, H2O et non pas une
atmosphère oxydante. Depuis l'expérience a été refaite plusieurs fois, en variant la composition de
l'atmosphère et la source d'énergie (utilisation du rayonnement ultraviolet notamment). Cependant, une
atmosphère moins réductrice (dioxyde de carbone CO2, azote N2, eau H2O) qui provient du volcanisme
donne de très mauvais rendements.
L'exploitation de l'idée de Miller [modifier]
Suite aux expériences de Miller, il a fallu déterminer les réactions chimiques qui se sont produites
dans l'enceinte (le ballon dans lequel il avait enfermé les différents gaz). Ainsi est née la chimie organique
dans l'eau.
Ces réactions nécessitent de fortes concentrations, des domaines de température et de pH très
étroits qui font que ces mécanismes sont très peu probables : une mare en voie d'assèchement pourrait
peut-être expliquer les fortes concentrations.
L'expérience fut à l'époque très critiquée à cause de cela. De plus, sa fiabilité a été remise en cause
car les molécules organiques obtenues pourraient a priori provenir d'une contamination extérieure. (?) Une
contamination extérieure est toutefois improbable pour les raisons évoquées ci-après.
En outre, la présence d'hydrogène (libre) dans l'atmosphère primitive est possible, mais seulement
en concentration réduite (de l'ordre du 1/100 de % ; ~100ppm ?), compte tenu de la masse de la Terre et de
la force de gravitation qui en résulte retenant très mal l'hydrogène vis à vis du vent solaire. Une des sources
d'hydrogène (libre) primitive aurait pu être les gaz volcaniques rejetant l'hydrogène issu de la réaction de
l'eau des roches avec des composés réducteurs.

L'expérience de Powner-Sutherland et l'origine des nucléotides à base pyrimidique

En 2009, Sutherland et son équipe[1] sont parvenu à établir une voie de synthèse des nucléotides à
base pyrimidique, l'acide uridylique et l'acide cytidylique. Les chercheurs butaient sur cette voie de synthèse
pré-biotique depuis 40 ans en raison de la difficulté de trouver la bonne façon de lier le ribose à la base
azotée. La clé de cette voie est de passer par un précurseur commun au ribose et la base azotée. Ce
précurseur, le 2-aminooxazol, est obtenu à partir de molécules organiques élémentaires : le glycéraldéhyde,
le cyanimide, le cyanoacétaldéhyde, le cyanoacétylène et le phosphate inorganique. Le mélange réactionnel
alimenté par de l'azote gazeux est soumis à un cycle de chauffage-refroidissement afin de simuler le cycle
d'évaporation d'une mare par le rayonnement solaire et l'alimentation par la pluie. Après une semaine le 2-
aminooxazol s'accumule dans le réacteur. Le précurseur se transforme ensuite en ribose et cytosine liés
ensemble. Le phosphate est ensuite ajouté au milieu réactionnel en présence d'UV durant trois jours
(l'absence d'ozone dans les conditions pré-biotiques engendrait un bombardement intense d'UV). L'acide
cytidylique se synthétise sous l'effet du rayonnement UV et quelques nucléotides portant une cytosine se
transforme en acide uridylique. Pour les deux autres nucléotides, l'équipe de Sutherland travaillent sur un
précurseur commun aux acides nucléiques à base purique.

Asymétrie des biomolécules [modifier]

L'Alanine, un acide aminé existant sous une forme lévogyre (L-Ala) ou dextrogyre (D-Ala).
Les molécules « chirales » sont des molécules pouvant exister sous deux formes possibles :
lévogyre (gauche) et dextrogyre (droite), de la même façon que la main gauche et la main droite sont l'image
symétrique l'une de l'autre. On appelle énantiomères ces différentes formes.
Une analyse poussée des molécules obtenues dans l'expérience de Miller montre que l'on obtient un
mélange racémique de molécules (autant de formes droites que de gauches), alors que l'on sait depuis le
milieu du XIXe siècle (notamment avec des travaux de Pasteur en 1847) que les acides aminés naturels
n'existent pratiquement que sous une de leurs deux formes énantiomères (on parle alors d'homochiralité) : la
forme lévogyre.
On a cependant retrouvé des traces d'acides aminés dextrogyres sous forme libre, dans des
peptides ou même des protéines. Ces formes, peu fréquentes, auraient par ailleurs des fonctions
physiologiques. Dans le monde vivant, on constate cependant que les sucres présents dans l'ADN sont
uniquement de type dextrogyre, ou que les agents de saveur ont un goût différent selon leur forme.

Théories sur l'origine de l'homochiralité [modifier]

Il existe deux grandes catégories de théories expliquant l'homochiralité : les théories biotiques et les
théories abiotiques.
Dans la première, on postule que la vie serait apparue à partir d'un mélange d'énantiomères, et que
l'homochiralité ne serait apparue que progressivement au cours de l'évolution. Selon Laurent Nahon, ce type
de théorie n'est cependant plus beaucoup soutenu, du fait que l'on a découvert que les protéines ne peuvent
se replier correctement si les acides aminés qui les composent ne sont pas chiraux. On estimerait donc que
l'homochiralité serait plutôt antérieure à l'apparition de la vie, ce sont les théories abiotiques.

Cristaux et énantiomères [modifier]

Les propriétés des cristaux permettent d'imaginer un scénario : certains cristaux inorganiques
exposent des faces possédant une chiralité intrinsèque, comme la calcite ou le quartz. Ces faces pourraient
adsorber préférentiellement une des deux formes énantiomères et donc la concentrer sélectivement aux
dépens de l'autre[2]. Cependant, rien n'indique que les cristaux aient pu jouer un rôle dans la formation de
molécules organiques.

Asymétricité et thermodynamique [modifier]

Il a été découvert que la matière est intrinsèquement asymétrique. « Lorsque l'on place des atomes
de cobalt dans une géométrie asymétrique, c'est-à-dire dans des champs magnétiques, les électrons
produits de la désintégration de neutron se déplacent toujours dans la direction opposée à leur spin (aligné
sur le champ). Les électrons sont donc intrinsèquement gauches. » Cette expérience de Tsung Dao Lee et
Chen Ning Yang qui reçurent le prix Nobel en 1957 peut être reproduite avec n'importe quel atome. Un gaz
de vapeur de césium par exemple dans un champ électromagnétique a un pouvoir rotatoire. C'est ce sur
quoi a travaillé Marie-Anne Bouchiat, directrice de recherche CNRS à l'ENS. Ce phénomène est facilement
observable sur les atomes lourds car la force mise en jeu est la force d'interaction faible entre le noyau et
Ainsi il a été calculé que les acides aminés naturels sont thermodynamiquement plus stables que
leur image dans un miroir.
Formation d'énantiomères dans l'espace [modifier]

Une météorite
Dans les années 1970, des acides aminés ont été découverts dans la météorite de Murchison, or, ils
étaient présents majoritairement sous leur forme lévogyre. On a ainsi découvert 70 acides aminés différents,
dont 3 seulement font partie des 20 acides aminés naturels. Le pourcentage exact de molécules lévogyres
est cependant beaucoup discuté, du fait d'éventuelles contaminations, et varie entre 50 % et -5 % suivant les
équipes de recherche.
L'idée que l'homochiralité aurait pour origine des molécules venues de l'espace s'est donc
Les énantiomères absorbent différemment la lumière lorsque celle-ci est polarisée «
circulairement » droite ou gauche. Or, ces molécules se dégradent après absorption, conduisant donc à un
excès d'une forme énantiomérique.

La nébuleuse d'Orion
On a découvert, en 1997, que la nébuleuse d'Orion produit de la lumière polarisée circulaire à 17 %
dans l'infrarouge (IR). L'infrarouge n'a pas assez d'énergie pour casser des liaisons covalentes, mais on peut
supposer que les ultraviolets (UV) sont également polarisés circulairement.
Une expérience menée en 2005 a montré qu'un mélange racémique d’un acide aminé simple irradié
par un rayonnement UV conduit à un mélange homochiral. Dans cette expérience, la leucine est utilisée à
l'état solide, reproduisant les conditions spatiales. Après une irradiation par un rayonnement synchrotron
polarisé circulairement droit (proche de la longueur d'onde observée dans l'espace, dans l'UV lointain),
l'expérience permet d'obtenir un excès de l'énantiomère lévogyre de 2,6 %.
Or, il suffit d'un excès de 1 % dans des réactions qui s'entretiennent pour conduire à un mélange
homochiral de 100 %. D'après Laurent Nahon, aucune expérience liée à des théories concurrentes n'est
parvenue à un tel excès.

Des molécules organiques aux protocellules [modifier]

Schéma d'une cellule animale.

Aujourd'hui, de nombreux modèles résolvent le problème de l'apparition des molécules organiques.
Les scientifiques arrivent à produire de nombreuses petites molécules biologiques (acides aminés, sucres,
bases nucléiques) dans des conditions prébiotiques en laboratoires.
Les expériences de Miller et les modèles qui en sont dérivés ne fournissent pas d'explication sur les
étapes suivantes qui incluent la transition des monomères aux biopolymères, puis aux protocellules et
finalement aux cellules vivantes ayant un métabolisme de base. Aussi les scientifiques ont exploré d'autres
voies de recherche.
Compartiments isolés [modifier]
L'apparition de compartiments isolés par une membrane pose de sérieux problèmes. Les
membranes des cellules vivantes sont composées de lipides, or on connaît aujourd'hui des acides gras à
longue chaîne qui peuvent spontanément former des petites membranes sphériques. Bien que l'on puisse
produire de tels compartiments en laboratoires, ces acides gras restent synthétisés par des enzymes. Le
processus permettant de former de tels compartiments en l'absence de ces enzymes demeure inconnu.

Protocellules [modifier]
Un compartiment isolé par une membrane ne forme cependant pas une protocellule. Selon Maynard
Smith, deux conditions sont nécessaires pour former une véritable protocellule :
1. Les molécules capables de répliquer la forme de base (les réplicateurs) doivent se lier entre
elles en un « chromosome », formant ainsi une unité structurelle, garantissant aux réplicateurs de
former un tout cohérent après la réplication ;
2. la membrane doit posséder des mécanismes d'échange avec le milieu extérieur, autres que
les systèmes à protéines actuels.
Schéma général d'un chromosome eucaryote. Schéma d'une membrane semi-perméable.
L'apparition du génome [modifier]

Les modèles « gènes d'abord » [modifier]

Une molécule d'ADN.

Dans ce modèle, l'apparition du génome a précédé l'apparition du métabolisme. Des molécules
d'ADN ou d'ARN auraient ainsi existé seules, s'autorépliquant à partir des molécules présentes dans leur
environnement. Les « individus » sont donc représentés par les molécules d'acides nucléiques elles-mêmes.
Les modèles « métabolisme d'abord » [modifier]
Plusieurs modèles rejettent l'idée de l'autoréplication d'un gène « nu » et font l'hypothèse de
l'apparition d'un métabolisme primitif qui aurait précédé l'émergence de la réplication de l'ARN. Une des
premières versions de cette hypothèse fut présentée en 1924 par Alexander Oparin avec son idée de
vésicules primitives capables de se répliquer, à une époque où on ne connaissait pas encore la structure de
D'autres variantes sont apparues dans les années 1980 et 1990 comme la théorie de Günter
Wächtershäuser sur un monde sulfuro-ferreux, ou les modèles de Christian de Duve basés sur la chimie des
D'autres arguments plus abstraits ont aussi été présentés. On peut citer les modèles mathématiques
de Freeman Dyson au début des années 1980 sur la probabilité de l'émergence d'un métabolisme sans
présence de gènes, ou encore les travaux de Stuart Kauffman sur les ensembles globalement
autocatalytiques (voir génération spontanée pour une présentation des idées de Kauffman sur l'origine de la

Métabolisme et génome : un monde à ARN [modifier]

Article détaillé : Hypothèse du monde à ARN.
L'hypothèse du monde à acide ribonucléique (ARN) est que l'ARN était la principale — et sans doute
la seule — forme de vie avant l'émergence de la première cellule à ADN. C'est Walter Gilbert qui a utilisé pour
la première fois le terme « monde à ARN » (« RNA world » en anglais) en 1986.
L'hypothèse d'un monde à ARN a aujourd'hui la faveur des scientifiques et est fondée sur plusieurs
éléments. Notamment sur le fait que l'ARN est en théorie capable aussi bien d'assurer des tâches
métaboliques que d'être le support d'une information génétique.

Stockage et réplication [modifier]

Une protéine liée à un brin d'ADN.

L'ARN a la capacité de stocker une information, en utilisant un code génétique similaire à celui de
l'ADN. L'ARN peut également se comporter comme un ribozyme (de la contraction de ribose et enzyme) et
catalyser certaines réactions, tout comme les protéines. Du point de vue de la reproduction, cette molécule
possède donc deux fonctions primordiales : le stockage de l'information et la catalyse nécessaire à
L'ADN peut aussi se recopier lui-même, mais seulement avec l'assistance de protéines. Les
protéines sont de très bons catalyseurs mais elles sont incapables de stocker l'information requise pour leur
propre réplication. L'ARN est lui capable à la fois de catalyse et d'autoréplication. Ainsi, le ribosome est un
ribozyme, dans le sens où le responsable de la synthèse des protéines n'est pas une protéine (comme c'est
le cas dans la grande majorité des catalyses d'une cellule vivante) mais l'ARN ribosomique lui-même. Ces
ribozymes peuvent se replier dans l'espace, faisant apparaître un site actif pour une catalyse, à l'instar des
L'ADN, formant une double hélice rigide, ne peut se replier pour jouer un rôle de catalyseur.

Efficacité des protéines [modifier]

La structure « quaternaire » d'une protéine.
Les protéines sont des catalyseurs très efficaces, bien plus que les ribozymes. De même, il existe 20
acides aminés dans le monde vivant, mais seulement quatre nucléotides, les protéines sont donc bien plus
diversifiées que les ARN.
D'un point de vue évolutif, il est donc peu probable qu'une protéine-enzyme ait été remplacée par
une ARN-enzyme. À l'inverse, si les ARN sont bien apparus avant les protéines, il est plausible qu'ils aient
été remplacés par des protéines, plus efficaces.
Cet argument est étayé par le fait que l'ARN joue un rôle dans la synthèse des protéines, via son rôle
fondamental dans les ribosomes actuels. L'ARN aurait donc en quelque sorte conduit à l'apparition des
Les protéines utilisées dans la structure du ribosome seraient donc venues plus tard : les premières
protéines auraient été sélectionnées selon leur capacité d'amélioration du fonctionnement des ribozymes,
pour finalement se substituer à eux.

Une grande distribution phylogénétique [modifier]

Les ARN sont présents dans les trois lignées du monde vivant (archées, procaryotes, eucaryotes).
Ils accomplissent dans chacun un grand nombre de tâches différentes, les plus connus sont l'ARN messager
(ARNm, véhiculant l'information génétique de l'ADN vers les ribosomes), l'ARN de transfert (ARNt, faisant le
lien entre acide nucléique et acide aminé) et l'ARN ribosomique (ARNr, composant structuraux et
fonctionnels des ribosomes). À côté de ceux-ci, on peut trouver un grand nombre d'ARN impliqué dans des
fonctions tels que des catalyses, des régulations de l'expression de gènes, des contrôles, des défenses
antivirales, des extinctions de gènes, des inhibitions de synthèses de protéines, des restaurations
génomiques, etc. C'est le cas des ARN interférents (ARNs), dont certains chercheurs qualifient le
mécanisme d'« universel ». Les ARNtm des procaryotes, ont également plusieurs fonctions : ils jouent à la
fois les rôles d'ARN de transfert et d'ARN messager.
Malgré cette grande diversité structurelle et fonctionnelle, la répartition des ARN permet de retrouver
le découpage du vivant. Ainsi, les petits ARN nucléolaire ne sont partagés que par les archées et les
eucaryotes, l'ARN de la télomérase n'est lui présent que chez les eucaryotes alors que les procaryotes sont
les seules à posséder des ARNtm. De la même façon, les trois grands types d'ARN (ARNt, ARNm et ARNr)
sont présents dans les trois lignées.
Le cas de l'ARN de transfert [modifier]
La structure de l'ARN de transfert.
Le rôle de l'ARNt est de transporter un acide aminé vers le ribosome, où s'effectuera la liaison avec
un autre acide aminé, pour former un polypeptide (donnant ainsi une protéine). Il existe plusieurs ARNt,
possédant chacun trois nucléotides : l'anticodon. L'anticodon correspond à un codon, porté par l'ARNm qui
définit l'ordre d'assemblage des acides aminés par le ribosome.
La particularité de l'ARNt est qu'il est, malgré sa petite taille, en partie constitué de nombreux
nucléotides que l'on ne rencontre pas ailleurs. Ces nucléotides « exotiques » auraient ainsi une origine
prébiotique, vestiges d'un monde à ARN. On retrouve ainsi ces composants dans l'ensemble des trois
domaines du vivant.

Virus et ARNt [modifier]

Il est relativement fréquent d'observer des virus à ARN ou des viroïdes portant des motifs similaires à
de l'ARNt. Ainsi, dans le viroïde PSTV (Potato Spindle Tuber Viroïd) et l'ARNt de la tyrosine, on trouve les
mêmes motifs structuraux en « feuille de trèfle ». Pour Marie-Christine Maurel, « ces derniers jouent un rôle
fondamental dans le vivant et leur ancienneté ne fait pas de doute ».
Autre structure surprenante : chez le virus TYMV (Turnip Yellow Mosaic Virus), l'amorce de la
traduction du génome du virus en protéine se fait par le biais d'une structure de type ARNt qui amorce sa
propre traduction et qui fixe un acide aminé.
Structure du virus PSTV.

Évolution dirigée d'ARN [modifier]

En 1990, Larry Gold et Jack Szostak ont mis au point une méthode visant à diriger l'évolution d'ARN,
afin de sélectionner ceux montrant une activité catalytique. Ils ont depuis réussi à obtenir des ribozymes
capables de lier des nucléotides entre eux, de lier des acides aminés à des ARNs, d'effectuer des réactions
d'oxydo-réductions, de se lier à des composants de la membrane, etc.
Il est donc en théorie possible, sur ce modèle, que l'ARN seul suffise à établir un métabolisme
primitif. Toutefois, il reste encore à découvrir un ARN capable de se répliquer lui-même.

ARN et hérédité [modifier]

L'ARN joue un rôle dans la transmission de l'activité des gènes : un tel mécanisme (qualifié
d'épigénétique) n'est pas lié à l'ADN, et serait une preuve de la capacité de l'ARN à participer à l'hérédité.

À l'origine de l'ADN dans la cellule [modifier]

Du point de vue de la biologie cellulaire, l'ADN est produit par modification d'un ARN : les
désoxyribonucléotides (précurseurs de l'ADN) sont en effet produits à partir des ribonucléotides (précurseurs
des ARN). De plus, le groupement thymine (identifié comme T dans le code génétique), est construit à partir
d'un groupement uracile (U). Or, bien que l'uracile soit spécifique à l'ARN et la thymine à l'ADN, le
groupement U est, au moment de la synthèse, déjà fixé sur un désoxyribonucléotide.
Uracile. Thymine.
De l'ARN à l'ADN [modifier]

Comparaison entre une molécule d'ARN (à gauche) et une molécule d'ADN (à droite).
Dans l'hypothèse du monde à ARN, apparaissent tout d'abord des viroïdes ressemblant à des ARN
auto-catalytiques, présents dans des compartiments isolés (qu'ils soient membranaires ou cristallins). Puis
des protocellules, capables de métabolisme archaïque, sont soumises à une évolution darwinienne, évoluant
ainsi vers des cellules à ARN, capables de présenter une activité variée et complexe.
Ribozymes ou protéines ? [modifier]
Bien que l'ARN soit donc à l'origine de l'ADN dans le métabolisme cellulaire, cette réaction est très
difficile à réaliser. De fait, dans les trois lignées, elle est catalysée par des protéines spécialisées : les
ribonucléotides-réductases. De plus, cette réaction est très coûteuse en énergie, du fait de la réduction du
ribose, et elle produit des radicaux libres, très réactifs, sur la protéine. L'ARN étant une molécule fragile, il
paraît improbable qu'elle puisse supporter des radicaux libres sans l'intervention de protéines.
Ainsi, l'origine de l'ADN trouve vraisemblablement sa source après l'apparition des protéines,
indispensables à chaque étape de sa synthèse à partir de précurseurs de type ARN, au sein de la cellule.
Intérêt de l'ADN [modifier]

Structure chimique de l'ADN.

L'ADN présente un certain nombre d'avantages sur l'ARN, en termes de conservation de l'intégrité
de l'information génétique.
Tout d'abord il se casse moins facilement, car le désoxyribose de l'ADN contient un atome d'oxygène
de moins que le ribose de l'ARN. Or l'oxygène peut facilement interagir sur les liaisons entre nucléotides,
posant alors un problème de stabilité.
Ensuite, l'ADN permet la réparation d'un problème récurrent : la transformation du groupement
thymine (T) en uracile (U). En effet, l'uracile, uniquement présent dans l'ARN, est une anomalie dans l'ADN.

Avantage sélectif de l'ADN : l'hypothèse du virus [modifier]

Structure de base d'un virus.

Les avantages en termes de stabilité de l'ADN pourraient ne pas suffire à expliquer son adoption.
Ainsi, Patrick Forterre avance l'hypothèse qu'un avantage sélectif supplémentaire peut être dû aux conflits
entre virus et cellules vivantes.
Dans ce modèle, le premier organisme à ADN serait un virus. L'ADN conférerait au virus le pouvoir
de résister à des enzymes dégradant les génomes à ARN, arme de défense probable des cellules. On
retrouve le même principe chez des virus actuels, qui altèrent leur ADN pour résister à des enzymes
produites par des bactéries infectées.
Actuellement, on peut observer que les enzymes nécessaires à la traduction de l'ARN vers l'ADN
sont très présentes chez les rétrovirus, dont le génome est porté par de l'ARN. De la même façon, de
nombreux virus codent leurs propres enzymes de synthèse de l'ADN.
Cette hypothèse est également corroborée par la découverte de virus à ADN, dont celui-ci contient,
non pas des groupements thymines, mais des groupements uraciles. Du point de vue évolutif, il y aurait donc
eu d'abord apparition des désoxyribonucléotides, puis de l'ADN à uracile (ADN-U), puis d'ADN à thymine
(ADN-T), qui se serait progressivement imposé. D'après Patrick Forterre, il est même probable que l'ADN-T
ait été « inventé deux fois », chez des virus différents.
Les virus à ARN seraient ici des reliques du monde à ARN, les virus à ADN-U seraient alors des
reliques du monde ayant précédé celui à ADN-T.
Les virus, premiers organismes à ADN [modifier]

Schéma de la réplication de l'ADN chez les eucaryotes.

Les virus à ADN pourraient être plus anciens que la première cellule à ADN : la première cellule à
ADN l'aurait donc emprunté à un ou plusieurs virus, sous la pression d'une course aux armements (théorie
de la reine rouge).
Didier Raoult et Jean-Michel Claverie ont ainsi découvert le mimivirus : un virus géant à ADN (son
génome étant deux fois plus long que le plus petit génome bactérien connu). La particularité de ce virus est
qu'il peut produire des protéines impliquées dans la traduction de l'ARN en protéines (comme des enzymes
chargeant des acides aminés sur des ARNt), il pourrait donc avoir pour ancêtres des virus plus anciens que
la première cellule à ADN.
Eugène Koonin et ses collègues ont mis en avant, en comparant des génomes séquencés, que la
plupart des enzymes impliqués dans la réplication de l'ADN sont différents entre les eubactéries et les
eucaryotes (accompagnés des archées). Ils en concluent que l'ADN aurait été inventé indépendamment
dans la lignée des eubactéries et celle conduisant aux eucaryotes et aux archées.[réf. nécessaire]
De même, les enzymes de réplications des virus à ADN sont très différentes d'un virus à l'autre, ainsi
que par rapport aux enzymes cellulaires jouant le même rôle.
Ces indices laissent penser que les enzymes liées à l'ADN sont apparues au cours d'un « premier
âge » du monde à ADN, où existaient cellules à ARN et virus à ARN et à ADN.

Passage de l'ADN dans les cellules [modifier]

La nature du génome du plus ancien ancêtre commun à tous les êtres vivants (que les scientifiques
prénomment LUCA) reste inconnue : faisait-il encore partie du monde à ARN, ou avait-il déjà un génome à
ADN ? Quoi qu'il en soit, LUCA est le fruit d'une longue évolution. Le génome des premières cellules, qui ont
précédé LUCA, était sans doute constitué par des molécules d'ARN et non pas d'ADN. Contrairement à
l'ADN, l'ARN peut en effet jouer à la fois le rôle d'enzyme et de matériel génétique. Stanley Miller et Christian
de Duve pensent que l'apparition de l'ARN a été elle-même un évènement tardif, en effet, cette molécule ne
semble pas pouvoir être synthétisée par les méthodes simples de la chimie prébiotique. L'ARN aurait donc
été précédé par des molécules dont nous ne connaîtrons sans doute jamais la nature exacte.
Ce qui amène les scientifiques à penser que l'ARN a précédé l'ADN résulte de ce constat : les trois
grandes lignées du vivant ne partagent que le système de synthèse des protéines, alors qu'elles diffèrent sur
le système de réplication de l'ADN.
Il existe alors plusieurs hypothèses pour le passage de l'ADN (d'origine viral) dans les cellules
vivantes : soit ce passage s'est produit une seule fois, soit il a pu avoir lieu plusieurs fois, indépendamment
dans les différentes lignées. Dans le premier cas, les premiers gènes des enzymes de réplications auraient
donc été remplacés par la suite par ceux d'un autre virus, formant ainsi les trois lignées.
L'hypothèse de Patrick Forterre est que les trois lignées du vivant trouvent leurs sources dans le
remplacement du génome à ARN par le génome à ADN de trois virus différents. On retrouve notamment ici
l'hypothèse d'une origine virale du noyau des eucaryotes.
Les travaux de Carl Woese semblent appuyer cette hypothèse, en démontrant que la vitesse
d'évolution des protéines semble avoir chuté au moment de l'apparition des trois lignées. Cette diminution
serait due au passage de l'ARN à l'ADN, les génomes à ADN étant plus stables, et donc moins sensibles aux
Dans ce scénario, l'apparition de trois lignées uniquement s'explique par le fait que les cellules à
ADN ont peu à peu supplanté les cellules à ARN, empêchant ainsi l'apparition de nouvelles lignées par
passage ARN→ADN.
Autres modèles [modifier]
Milieu de collision Planète/Noyaux de comètes [modifier]
Selon des travaux de modélisation travaux présentés en 2010 [3], la compression brutale des
matériaux constituant le noyau de glace d'une comète, au moment du choc avec une planète pourrait donner
lieu à la production d'acides aminés. Cette modélisation a pris comme base 210 molécules diluées dans un
mélange d'eau, de méthanol, d'ammoniac, de dioxyde de carbone et de monoxyde de carbone estimés
couramment présentes dans le noyau des comètes. Avec une vitesse d'impact de 29 km/s, la pression atteint
10 gigapascals et une température de 700°Kelvin. Selon les auteurs, d'autres simulations, à pressions et
températures plus élevées conduisent à des réactions chimiques encore plus complexes. Selon le modèle,
une simulation de 47 gigapascals et une température de 3.141 Kelvin pour les 20 premières picosecondes
de l'impact fait naitre des molécules complexes, dont de grosses molécules à liaisons carbone-azote qui
pourraient être les premières briques de la vie. Le calcul a nécessité environ million d'heures d'ordinateur sur
le superordinateur du cluster Atlas du Lawrence Livermore.

Sources hydrothermales : le monde du soufre [modifier]

Les sources hydrothermales ont été découvertes en 1977 à 2 600 mètres de profondeur, là où deux
plaques tectoniques se séparent.
Les monts hydrothermaux sont situés sur la couche sédimentaire. Leur diamètre à la base varie de
25 à 100 mètres et leur hauteur varie de 70 à 100 mètres. Les cheminées de ces fumeurs sont parfois
recouvertes d'une croûte d'oxyde de manganèse. Les fumeurs situés sur ces monts sont composés d'un
solide friable dont la couleur varie du gris noir à l'ocre, ce sont des sulfures de fer, de cuivre et de zinc.
Ces sources sont particulièrement intéressantes car on y a trouvé la vie où on la croyait impossible :
milieu privé d'oxygène, à haute température, chargé de métaux et de soufre, dans l'obscurité la plus totale.
Cependant les gradients de température importants autour de ces zones et le fait que les ultraviolets
destructeurs ne parviennent pas si profondément (alors qu'ils détruisent toute molécule formée à la surface)
sont de bonnes conditions pour l'apparition de la vie.
Ces organismes ont les mêmes formes que ceux que l'on connaît plus près de la surface (ADN,
protéines, sucres...) mais puisent leur énergie de l'oxydation du sulfure d'hydrogène (H2S) pour transformer
le carbone minéral en matière organique.
D'autre part, des expériences ont été menées, au laboratoire de géophysique de Washington, et ont
montré que dans les conditions qui existent autour des évents, il y a formation d'ammoniac (NH3), forme
réduite de l'azote qui est tant nécessaire à la formation des molécules organiques de la première partie et qui
n'existait pas dans l'atmosphère oxydante. Les sources hydrothermales sont donc de bonnes sources de

Une origine extraterrestre primitive (exogenèse) [modifier]

Une hypothèse alternative est que la vie se soit d'abord formée hors de la Terre. Les composés
organiques sont relativement fréquents dans l'espace, notamment dans les zones lointaines du système
solaire où l'évaporation des composés volatils est très réduite. Certaines comètes sont enrobées dans des
couches de matière sombre, qu'on pense être une sorte de bitume formé par une combinaison de composés
carbonés simples et de rayons ultraviolets. La pluie de matériaux cométaires sur la Terre primitive pourrait
avoir apporté des quantités de molécules organiques complexes, ce qui aurait causé l'apparition de la vie sur
Une hypothèse plus large est la panspermie : la vie même serait apparue dans l'espace puis
disséminée sur Terre. Selon une variante, la vie serait apparue sur Mars d'abord et des impacts de comètes
et d'astéroïdes sur Mars auraient projeté du matériel de la surface martienne sur Terre. Il est encore plus
difficile de trouver des indices pour justifier ces hypothèses que les théories plus classiques.
Ces théories d'une origine extraterrestre n'expliquent pas directement comment la vie est apparue,
car elles ne font a priori que reporter le problème. Cependant, elles élargissent les conditions dans
lesquelles la vie a pu apparaître dans l'univers. Les futurs échantillons de sols ramenés de Mars et de
comètes permettront peut-être d'obtenir de nouveaux éléments de réponse.

Notes et références [modifier]

1. ↑ Matthew W. Powner, Béatrice Gerland and John D. Sutherland, Synthesis of activated
pyrimidine ribonucleotides in prebiotically plausible conditions, Nature, Vol 459, 14 May 2009, pp.
2. ↑ Robert M. Hazen, Timothy R. Filley, and Glenn A. Goodfriend, PNAS May 8, 2001 vol. 98
no. 10 5487–5490Y
3. ↑ travaux de Nir Goldman et de ses collègues du Lawrence Livermore National Laboratory à
Livermore, présentés le 24 Mars 2010 devant l'American Chemical Society à San Francisco,
Californie, et relayés par une brève du Journal Nature [archive] (26 Mars 2010 ; On line :
Sources [modifier]
• (en) John Desmond Bernal, The origin of life, éditions Weidenfeld & Nicolson, 1967.
• (fr) Guillemette Lauters, Olivier Lenaerts, Daniel Rousselet et Eric Depiereux, Bioscope :
L’origine de la vie.
• (fr) John Maynard Smith, Eörs Szathmáry, Les Origines de la vie, éditions Dunod, 2000.
(ISBN 2100048600)
• (en) J. William Schopf, Anatoliy B. Kudryavtsev, David G. Agresti, Thomas J. Wdowiak,
Andrew D. Czaja, « Laser−Raman imagery of Earth's earliest fossils », dans Nature, volume 416,
pages 73–76, mars 2002 [(en) lire en ligne].
• (fr) Marie-Christine Maurel, La Naissance de la vie : de l'évolution prébiotique à
l'évolution biologique, éditions Dunod, 2003 (3e édition). (ISBN 2100068822)
• (en) N. Fujii, T. Saito, « Homochirality and life », dans The Chemical Records, volume 4,
n 5, pages 267–78, 2004 [(en) lire en ligne]
• (fr) Uwe Meierhenrich, Un pas vers l’origine de l’homochiralité…, novembre 2005.
• (fr) Patrick Forterre, « L'origine du génome », dans Les dossiers de La Recherche :
l'histoire de la vie, les grandes étapes de l'évolution , no 19, pages 34–40, mai 2005.
• (fr) Marc-André Selosse, « Quelle parenté entre les trois grandes lignées du vivant ? »,
dans Les dossiers de La Recherche : l'histoire de la vie, les grandes étapes de l'évolution , no 19,
pages 42–45, mai 2005.
• (fr) « Laurent Nahon : « L'asymétrie des biomolécules vient de l'espace » », propos
recueillis par Franck Daninos, dans La Recherche, no 390, octobre 2005.
• (en) U. Meierhenrich, L. Nahon, C. Alcaraz, J. Bredehöft, S. Hoffmann, B. Barbier, A.
Brack, « Asymmetric VUV photodecomposition of the amino acid D,L-Leucine in the solid state »,
dans Angewandte Chemie International Edition, volume 44, pages 5630–5634, juillet 2005 [(en) lire
en ligne].

Bibliographie [modifier]
• (fr) Le site de la Société Française d'Exobiologie (SFE).
• (fr) Conférence de Patrick Forterre : "Regards croisés sur les origines de la vie" (2009) à
l'ENS Cachan.
• (en) Uwe Meierhenrich: Amino acids and the asymmetry of life, Springer-Verlag, 2008.
(ISBN 978-3-540-76885-2)
• (en) Site dédié au mimivirus.
• (en) Anthony M. Poole, What is the Last Universal Common Ancestor ?, septembre 2002.
• (en) Antonio Lazcano The Origins of Life. Have too many cooks spoiled the prebiotic
soup?, Natural History magazine, février 2006.
• (en) Erwin Schrödinger, What is Life?, 1944. (fr) Traduction française : Qu'est-ce que
la vie?, ed. Seuil.
• (fr) Joël de Rosnay, Les Origines de la vie, éditions du Seuil, 1966. (ISBN 2020002167)
• (fr) Jean C. Baudet, La vie expliquée par la chimie, Vuibert, Paris, 2006.
• (fr) André Brack, Bénédicte Leclercq, La vie est-elle universelle ? – Des premiers êtres
vivants à l'exploration spatiale, éditions EDP Sciences, 2003. (ISBN 2-86883-674-7).
• (fr) Sous la direction de Muriel Gargaud, Didier Despois, Jean-Paul Parisot,
L'Environnement de la Terre Primitive, Collection : L'origine de la Vie sur Terre et dans l'Univers,
Presses universitaires de Bordeaux, 2001. (ISBN 2-86781-267-4).

Voir aussi [modifier]

Articles connexes [modifier]
• Biologie, Biologie cellulaire, vie
• Biochimies hypothétiques
• Exobiologie
• Habitabilité d'une planète
• Génération spontanée
• Protéinoïde

Liens externes [modifier]

• (fr) Écouter La leçon consacrée aux origines chimiques de la vie et dispensée par
Jacques Reisse au Collège Belgique (2009).
• (fr) Comprendre facilement l'origine de la vie sur la Terre
• (fr) Un site qui explique avec des mots simples les origines de la vie
• (fr) Les trois grandes théories de l'apparition de la vie sur Terre

Évolution biologique

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et


Dérive génétique · Mutation génétique · Recombinaison

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis
[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation
Branches de la biologie
Anatomie · Biochimie · Bio-informatique · Biologie cellulaire · Biologie de l'évolution · Biologie
humaine · Biologie marine · Biologie moléculaire · Botanique · Écologie · Exobiologie · Génétique ·
Génomique · Géobiologie · Microbiologie · Origine de la vie · Paléontologie · Parasitologie · Physiologie ·
Taxinomie · Virologie · Zoologie

• Portail de la chimie

• Portail de la biochimie

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « ».

Catégories : Origine de la vie | Histoire évolutive | Métabolisme | [+]


From Wikipedia, the free encyclopedia

Jump to: navigation, search

For other uses, see Life (disambiguation).

Life (Biota)
Life on a rocky peak in the Waitakere Ranges

Scientific classification

Domains and Kingdoms

Life on Earth:
• Non-cellular life (viruses) * *
• Cellular life
• Bacteria
• Archaea
• Eukarya
• Protista
• Fungi
• Plantae
• Animalia
Life (cf. biota) is a characteristic that distinguishes objects that have signaling and self-sustaining
processes (biology) from those that do not,[1][2] either because such functions have ceased (death), or else
because they lack such functions and are classified as inanimate.[3]
In biology, the science of living organisms, life is the condition which distinguishes active organisms
from inorganic matter.[4] Living organisms undergo metabolism, maintain homeostasis, possess a capacity
to grow, respond to stimuli, reproduce and, through natural selection, adapt to their environment in
successive generations. More complex living organisms can communicate through various means.[1][5] A
diverse array of living organisms (life forms) can be found in the biosphere on Earth, and the properties
common to these organisms—plants, animals, fungi, protists, archaea, and bacteria—are a carbon- and water-
based cellular form with complex organization and heritable genetic information.
In philosophy and religion, the conception of life and its nature varies. Both offer interpretations as to
how life relates to existence and consciousness, and both touch on many related issues, including life
stance, purpose, conception of a god or gods, a soul or an afterlife.
• 1 Early theories about life
• 1.1 Materialism
• 1.2 Hylomorphism
• 1.3 Vitalism
• 2 Definitions
• 2.1 Biology
• 2.2 Biophysics
• 2.3 Living systems theories
• 3 Origin of life
• 4 Conditions for life
• 4.1 Range of tolerance
• 4.2 Extremophiles
• 5 Classification of life
• 6 Extraterrestrial life
• 7 Death
• 7.1 Extinction
• 7.2 Fossils
• 8 See also
• 9 References
[edit] Early theories about life
[edit] Materialism

Plant life
Herds of zebra and impala gathering on the Masai Mara plain

An aerial photo of microbial mats around the Grand Prismatic Spring of Yellowstone National Park.
Some of the earliest theories of life were materialist, holding that all that exists is matter, and that all
life is merely a complex form or arrangement of matter. Empedocles (430 B.C.) argued that every thing in the
universe is made up of a combination of four eternal 'elements' or 'roots of all': earth, water, air, and fire. All
change is explained by the arrangement and rearrangement of these four elements. The various forms of life
are caused by an appropriate mixture of elements. For example, growth in plants is explained by the natural
downward movement of earth and the natural upward movement of fire.[6]
Democritus (460 B.C.), the disciple of Leucippus, thought that the essential characteristic of life is
having a soul (psychê). In common with other ancient writers, he used the term to mean the principle of living
things that causes them to function as a living thing. He thought the soul was composed of fire atoms,
because of the apparent connection between life and heat, and because fire moves.[7] He also suggested
that humans originally lived like animals, gradually developing communities to help one another, originating
language, and developing crafts and agriculture.[8]
In the scientific revolution of the seventeenth century, mechanistic ideas were revived by
philosophers like Descartes.

[edit] Hylomorphism
Hylomorphism is the theory (originating with Aristotle (322 BC)) that all things are a combination of
matter and form. Aristotle was one of the first ancient writers to approach the subject of life in a scientific way.
Biology was one of his main interests, and there is extensive biological material in his extant writings.
According to him, all things in the material universe have both matter and form. The form of a living thing is its
soul (Greek 'psyche', Latin 'anima'). There are three kinds of souls: the 'vegetative soul' of plants, which
causes them to grow and decay and nourish themselves, but does not cause motion and sensation; the
'animal soul' which causes animals to move and feel; and the rational soul which is the source of
consciousness and reasoning which (Aristotle believed) is found only in man.[9] Each higher soul has all the
attributes of the lower one. Aristotle believed that while matter can exist without form, form cannot exist
without matter, and therefore the soul cannot exist without the body.[10]
Consistent with this account is a teleological explanation of life. A teleological explanation accounts
for phenomena in terms of their purpose or goal-directedness. Thus, the whiteness of the polar bear's coat is
explained by its purpose of camouflage. The direction of causality is the other way round from materialistic
science, which explains the consequence in terms of a prior cause. Modern biologists now reject this
functional view in terms of a material and causal one: biological features are to be explained not by looking
forward to future optimal results, but by looking backwards to the past evolutionary history of a species,
which led to the natural selection of the features in question.

[edit] Vitalism
Vitalism is the belief that the life-principle is essentially immaterial. This originated with Stahl (17th
century), and held sway until the middle of the nineteenth century. It appealed to philosophers such as Henri
Bergson, Nietzsche, Wilhelm Dilthey, anatomists like Bichat, and chemists like Liebig.
Vitalism underpinned the idea of a fundamental separation of organic and inorganic material, and the
belief that organic material can only be derived from living things. This was disproved in 1828 when Wöhler
prepared urea from inorganic materials. This so-called Wöhler synthesis is considered the starting point of
modern organic chemistry. It is of great historical significance because for the first time an organic compound
was produced from inorganic reactants.
Later, Helmholtz, anticipated by Mayer, demonstrated that no energy is lost in muscle movement,
suggesting that there were no vital forces necessary to move a muscle. These empirical results led to the
abandonment of scientific interest in vitalistic theories, although the belief lingered on in non-scientific
theories such as homeopathy, which interprets diseases and sickness as caused by disturbances in a
hypothetical vital force or life force.

[edit] Definitions
It is still a challenge for scientists and philosophers to define life in unequivocal terms.[11][12][13]
Defining life is difficult —in part— because life is a process, not a pure substance.[14] Any definition must be
sufficiently broad to encompass all life with which we are familiar, and it should be sufficiently general that,
with it, scientists would not miss life that may be fundamentally different from earthly life.[15]

[edit] Biology
Since there is no unequivocal definition of life, the current understanding is descriptive, where life is a
characteristic of organisms that exhibit all or most of the following phenomena:[14][16][17]
1. Homeostasis: Regulation of the internal environment to maintain a constant state; for
example, electrolyte concentration or sweating to reduce temperature.
2. Organization: Being structurally composed of one or more cells, which are the basic units of
3. Metabolism: Transformation of energy by converting chemicals and energy into cellular
components (anabolism) and decomposing organic matter (catabolism). Living things require energy
to maintain internal organization (homeostasis) and to produce the other phenomena associated with
4. Growth: Maintenance of a higher rate of anabolism than catabolism. A growing organism
increases in size in all of its parts, rather than simply accumulating matter.
5. Adaptation: The ability to change over a period of time in response to the environment. This
ability is fundamental to the process of evolution and is determined by the organism's heredity as well
as the composition of metabolized substances, and external factors present.
6. Response to stimuli: A response can take many forms, from the contraction of a unicellular
organism to external chemicals, to complex reactions involving all the senses of multicellular
organisms. A response is often expressed by motion, for example, the leaves of a plant turning
toward the sun (phototropism) and by chemotaxis.
7. Reproduction: The ability to produce new individual organisms, either asexually from a single
parent organism, or sexually from two parent organisms.
To reflect the minimum phenomena required, some have proposed other biological definitions of life:
• Living things are systems that tend to respond to changes in their environment, and inside
themselves, in such a way as to promote their own continuation.[17]
• A network of inferior negative feedbacks (regulatory mechanisms) subordinated to a superior
positive feedback (potential of expansion, reproduction).[18]
• A systemic definition of life is that living things are self-organizing and autopoietic (self-
producing). Variations of this definition include Stuart Kauffman's definition as an autonomous agent
or a multi-agent system capable of reproducing itself or themselves, and of completing at least one
thermodynamic work cycle.[19]
• Life is a self-sustained chemical system capable of undergoing Darwinian evolution.[20]
• Things with the capacity for metabolism and motion.[14]
Viruses are most often considered replicators rather than forms of life. They have been described as
"organisms at the edge of life",[21] since they possess genes, evolve by natural selection,[22] and replicate
by creating multiple copies of themselves through self-assembly. However, viruses do not metabolize and
require a host cell to make new products. Virus self-assembly within host cells has implications for the study
of the origin of life, as it may support the hypothesis that life could have started as self-assembling organic

[edit] Biophysics
Biophysicists have also commented on the nature and qualities of life forms—notably that they
function on negative entropy.[25][26] In more detail, according to physicists such as John Bernal, Erwin
Schrödinger, Eugene Wigner, and John Avery, life is a member of the class of phenomena which are open or
continuous systems able to decrease their internal entropy at the expense of substances or free energy taken
in from the environment and subsequently rejected in a degraded form (see: entropy and life).[27][28][29]

[edit] Living systems theories

Some scientists have proposed in the last few decades that a general living systems theory is
required to explain the nature of life.[30] Such general theory, arising out of the ecological and biological
sciences, attempts to map general principles for how all living systems work. Instead of examining
phenomena by attempting to break things down into component parts, a general living systems theory
explores phenomena in terms of dynamic patterns of the relationships of organisms with their environment.
Gaia hypothesis
The idea that the Earth is alive is probably as old as humankind, but the first public expression of it as
a fact of science was by a Scottish scientist, James Hutton. In 1785 he stated that the Earth was a
superorganism and that its proper study should be physiology. Hutton is rightly remembered as the father of
geology, but his idea of a living Earth was forgotten in the intense reductionism of the nineteenth century.[32]
The Gaia hypothesis, originally proposed in the 1960s by scientist James Lovelock,[33][34] explores the idea
that the life on Earth functions as a single organism which actually defines and maintains environmental
conditions necessary for its survival.[35]
Robert Rosen (1991) built on the assumption that the explanatory powers of the mechanistic
worldview cannot help understand the realm of living systems. One of several important clarifications he
made was to define a system component as "a unit of organization; a part with a function, i.e., a definite
relation between part and whole." From this and other starting concepts, he developed a "relational theory of
systems" that attempts to explain the special properties of life. Specifically, he identified the
"nonfractionability of components in an organism" as the fundamental difference between living systems and
'biological machines.'[36]
Life as a property of ecosystems
A systems view of life treats environmental fluxes and biological fluxes together as a "reciprocity of
influence",[37] and a reciprocal relation with environment is arguably as important for understanding life as it
is for understanding ecosystems. As Harold J. Morowitz (1992) explains it, life is a property of an ecological
system rather than a single organism or species.[38] He argues that an ecosystemic definition of life is
preferable to a strictly biochemical or physical one. Robert Ulanowicz (2009) also highlights mutualism as the
key to understand the systemic, order-generating behavior of life and ecosystems.[39]

[edit] Origin of life

Main article: Origin of life
For religious beliefs about the creation of life, see creation myth.
Evidence suggests that life on Earth has existed for about 3.7 billion years.[40] All known life forms
share fundamental molecular mechanisms, and based on these observations, theories on the origin of life
attempt to find a mechanism explaining the formation of a primordial single cell organism from which all life
originates. There are many different hypotheses regarding the path that might have been taken from simple
organic molecules via pre-cellular life to protocells and metabolism. Many models fall into the "genes-first"
category or the "metabolism-first" category, but a recent trend is the emergence of hybrid models that
combine both categories.[41]
There is no scientific consensus as to how life originated and all proposed theories are highly
speculative. However, most currently accepted scientific models build in one way or another on the following
• The Miller-Urey experiment, and the work of Sidney Fox, suggest that that conditions on the
primitive Earth may have favored chemical reactions that synthesized some amino acids and other
organic compounds from inorganic precursors.
• Phospholipids spontaneously form lipid bilayers, the basic structure of a cell membrane.
Life as we know it today synthesizes proteins, which are polymers of amino acids using instructions
encoded by cellular genes—which are polymers of deoxyribonucleic acid (DNA). Protein synthesis also
entails intermediary ribonucleic acid (RNA) polymers. One possibility is that genes came first[42] and then
proteins. Another possibility is that proteins came first[43] and then genes. However, because genes are
required to make proteins, and proteins are required to make genes, the problem of considering which came
first is like that of the chicken or the egg. Most scientists have adopted the hypothesis that because DNA and
proteins function together so intimately, it's unlikely that they arose independently.[44] Therefore, many
scientists consider the possibility, apparently first suggested by Francis Crick,[45] that the first life was based
on the DNA-protein intermediary: RNA.[44] In fact, RNA has the DNA-like properties of information storage
and replication and the catalytic properties of some proteins. Crick and others actually favored the RNA-first
hypothesis[46] even before the catalytic properties of RNA had been demonstrated by Thomas Cech.[47]
A significant issue with the RNA-first hypothesis is that experiments designed to synthesize RNA
from simple precursors have not been nearly as successful as the Miller-Urey experiments that synthesized
other organic molecules from inorganic precursors. One reason for the failure to create RNA in the laboratory
is that RNA precursors are very stable and don't react with each other under ambient conditions. However,
the successful synthesis of certain RNA molecules under conditions hypothesized to exist prior to life on
Earth has been achieved by adding alternative precursors in a specified order with the precursor phosphate
present throughout the reaction.[48] This study makes the RNA-first hypothesis more plausible to many
Recent experiments have demonstrated true Darwinian evolution of unique RNA enzymes
(ribozymes) made up of two separate catalytic components that replicate each other in vitro.[50] In describing
this work from his laboratory, Gerald Joyce stated: "This is the first example, outside of biology, of
evolutionary adaptation in a molecular genetic system."[51] Such experiments make the possibility of a
primordial RNA World even more attractive to many scientists.

[edit] Conditions for life

The diversity of life on Earth today is a result of the dynamic interplay between genetic opportunity,
metabolic capability, environmental challenges,[52] and symbiosis.[53][54][55] For most of its existence,
Earth's habitable environment has been dominated by microorganisms and subjected to their metabolism
and evolution. As a consequence of such microbial activities on a geologic time scale, the physical-chemical
environment on Earth has been changing, thereby determining the path of evolution of subsequent life.[52]
For example, the release of molecular oxygen by cyanobacteria as a by-product of photosynthesis induced
fundamental, global changes in the Earth's environment. The altered environment, in turn, posed novel
evolutionary challenges to the organisms present, which ultimately resulted in the formation of our planet's
major animal and plant species. Therefore this 'co-evolution' between organisms and their environment is
apparently an inherent feature of living systems.[52]

[edit] Range of tolerance

The inert components of an ecosystem are the physical and chemical factors necessary for life –
energy (sunlight or chemical energy), water, temperature, atmosphere, gravity, nutrients, and ultraviolet solar
radiation protection.[56] In most ecosystems the conditions vary during the day and often shift from one
season to the next. To live in most ecosystems, then, organisms must be able to survive a range of
conditions, called 'range of tolerance'.[57] Outside of that are the 'zones of physiological stress', where the
survival and reproduction are possible but not optimal. Outside of these zones are the 'zones of intolerance',
where life for that organism is implausible. It has been determined that organisms that have a wide range of
tolerance are more widely distributed than organisms with a narrow range of tolerance.[57]

[edit] Extremophiles

Deinococcus radiodurans can resist radiation exposure.

Main article: Extremophile
To survive, some microorganisms can assume forms that enable them to withstand freezing,
complete desiccation, starvation, high-levels of radiation exposure, and other physical or chemical
challenges. Furthermore, some microorganisms can survive exposure to such conditions for weeks, months,
years, or even centuries.[52] Extremophiles are microbial life forms that thrive outside the ranges life is
commonly found in. They also excel at exploiting uncommon sources of energy. While all organisms are
composed of nearly identical molecules, evolution has enabled such microbes to cope with this wide range of
physical and chemical conditions. Characterization of the structure and metabolic diversity of microbial
communities in such extreme environments is ongoing. An understanding of the tenacity and versatility of life
on Earth, as well as an understanding of the molecular systems that some organisms utilize to survive such
extremes, will provide a critical foundation for the search for life beyond Earth.[52]

[edit] Classification of life

Main article: Biological classification
The hierarchy of biological classification's eight major taxonomic ranks, which is an example of
definition by genus and differentia. Life is divided into domains, which are subdivided into further groups.
Intermediate minor rankings are not shown.
Traditionally, people have divided organisms into the classes of plants and animals, based mainly on
their ability of movement. The first known attempt to classify organisms was conducted by the Greek
philosopher Aristotle (384-322 BC). He classified all living organisms known at that time as either a plant or
an animal. Aristotle distinguished animals with blood from animals without blood (or at least without red
blood), which can be compared with the concepts of vertebrates and invertebrates respectively. He divided
the blooded animals into five groups: viviparous quadrupeds (mammals), birds, oviparous quadrupeds
(reptiles and amphibians), fishes and whales. The bloodless animals were also divided into five groups:
cephalopods, crustaceans, insects (which also included the spiders, scorpions, and centipedes, in addition to
what we now define as insects), shelled animals (such as most molluscs and echinoderms) and "zoophytes".
Though Aristotle's work in zoology was not without errors, it was the grandest biological synthesis of the time
and remained the ultimate authority for many centuries after his death.[58]
The exploration of the American continent revealed large numbers of new plants and animals that
needed descriptions and classification. In the latter part of the 16th century and the beginning of the 17th,
careful study of animals commenced and was gradually extended until it formed a sufficient body of
knowledge to serve as an anatomical basis for classification.
In the late 1740s, Carolus Linnaeus introduced his method, still used, to formulate the scientific name
of every species.[59] Linnaeus took every effort to improve the composition and reduce the length of the
many-worded names by abolishing unnecessary rhetoric, introducing new descriptive terms and defining
their meaning with an unprecedented precision. By consistently using his system, Linnaeus separated
nomenclature from taxonomy. This convention for naming species is referred to as binomial nomenclature.
The fungi were originally treated as plants. For a short period Linnaeus had placed them in the taxon
Vermes in Animalia. He later placed them back in Plantae. Copeland classified the Fungi in his Protoctista,
thus partially avoiding the problem but acknowledged their special status.[60] The problem was eventually
solved by Whittaker, when he gave them their own kingdom in his five-kingdom system. As it turned out, the
fungi are more closely related to animals than to plants.[61]
As new discoveries enabled us to study cells and microorganisms, new groups of life were revealed,
and the fields of cell biology and microbiology were created. These new organisms were originally described
separately in protozoa as animals and protophyta/thallophyta as plants, but were united by Haeckel in his
kingdom protista, later the group of prokaryotes were split off in the kingdom Monera, eventually this kingdom
would be divided in two separate groups, the Bacteria and the Archaea, leading to the six-kingdom system
and eventually to the current three-domain system.[62] The classification of eukaryotes is still controversial,
with protist taxonomy especially problematic.[63]
As microbiology, molecular biology and virology developed, non-cellular reproducing agents were
discovered, such as viruses and viroids. Sometimes these entities are considered to be alive but others
argue that viruses are not living organisms since they lack characteristics such as cell membrane,
metabolism and do not grow or respond to their environments. Viruses can however be classed into
"species" based on their biology and genetics but many aspects of such a classification remain controversial.
Since the 1960s a trend called cladistics has emerged, arranging taxa in an evolutionary or
phylogenetic tree. It is unclear, should this be implemented, how the different codes will coexist.[65]

Linnaeus Haeckel Chatton Copeland Whittaker Woese et al.

1735[66] 1866[67] 1925[68][69] 1938[60][70] 1969[71] 1977[72][73]
2 kingdoms 3 kingdoms 2 empires 4 kingdoms 5 kingdoms 6 kingdoms

Prokaryota Monera Monera
(not Archaebacteria
Protista Protista

Eukaryota Fungi Fungi

Vegetabilia Plantae
Plantae Plantae Plantae

Animalia Animalia Animalia Animalia Animalia

[edit] Extraterrestrial life
Main articles: Extraterrestrial life and astrobiology

Panspermia hypothesis
Earth is the only planet in the universe known to harbour life. The Drake equation, which relates the
number of extraterrestrial civilizations in our galaxy with which we might come in contact, has been used to
discuss the probability of life elsewhere, but scientists disagree on many of the values of variables in this
equation. Depending on those values, the equation may either suggest that life arises frequently or
The region around a main sequence star that could support Earth-like life on an Earth-like planet is
known as the habitable zone. The inner and outer radii of this zone vary with the luminosity of the star, as
does the time interval during which the zone will survive. Stars more massive than the Sun have a larger
habitable zone, but will remain on the main sequence for a shorter time interval during which life can evolve.
Small red dwarf stars have the opposite problem, compounded with higher levels of magnetic activity and the
effects of tidal locking from close orbits. Hence, stars in the intermediate mass range such as the Sun may
possess the optimal conditions for Earth-like life to develop. The location of the star within a galaxy may also
have an impact on the likelihood of life forming.
Panspermia, also called exogenesis, is a hypothesis proposing that life originated elsewhere in the
universe and was subsequently transferred to Earth in the form of spores perhaps via meteorites, comets or
cosmic dust. However, this hypothesis does not help explain the ultimate origin of life.

[edit] Death
Main article: Death
Death is the permanent termination of all vital functions or life processes in an organism or cell.[75]
[76] After death, the remains of an organism become part of the biogeochemical cycle. Organisms may be
consumed by a predator or a scavenger and leftover organic material may then be further decomposed by
detritivores, organisms which recycle detritus, returning it to the environment for reuse in the food chain.
One of the challenges in defining death is in distinguishing it from life. Death would seem to refer to either the
moment at which life ends, or when the state that follows life begins.[77] However, determining when death
has occurred requires drawing precise conceptual boundaries between life and death. This is problematic,
however, because there is little consensus over how to define life. The nature of death has for millennia been
a central concern of the world's religious traditions and of philosophical inquiry. Many religions maintain faith
in either some kind of afterlife, reincarnation, or resurrection.

[edit] Extinction
Main article: Extinction
Extinction is the gradual process by which a group of taxa or species dies out, reducing biodiversity.
[78] The moment of extinction is generally considered to be the death of the last individual of that species.
Because a species' potential range may be very large, determining this moment is difficult, and is usually
done retrospectively after a period of apparent absence. Species become extinct when they are no longer
able to survive in changing habitat or against superior competition. Over the history of the Earth, over 99% of
all the species that have ever lived have gone extinct,[79] however, mass extinctions may have accelerated
evolution by providing opportunities for new groups of organisms to diversify.[80]

[edit] Fossils
Main article: Fossil
Fossils are the preserved remains or traces of animals, plants, and other organisms from the remote
past. The totality of fossils, both discovered and undiscovered, and their placement in fossil-containing rock
formations and sedimentary layers (strata) is known as the fossil record. Such a preserved specimen is
called a "fossil" if it is older than the arbitrary date of 10,000 years ago.[81] Hence, fossils range in age from
the youngest at the start of the Holocene Epoch to the oldest from the Archaean Eon, a few billion years old.

[edit] See also

• Alpha taxonomy • Kingdom (biology)
• Artificial life • Life on Mars
• Biological immortality • Meaning of life
• Biology - the study of life • Nature
• Carbon-based life • Non-cellular life
• Cellular life • Organic life
• Conway's Game of Life • Organism
• Death • Origin of life
• Entropy and life • Personal life
• Evolutionary history of life • Phylogenetics
• Extraterrestrial life • Prehistoric life
• Extremophile • Prion
• Gaia hypothesis • Quality of life
• Genetics • Silicon-based life
• Genetic engineering • Synthetic life
• Hierarchy of life
• Hypothetical types of

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[edit] Further reading

• Kauffman, Stuart. The Adjacent Possible: A Talk with Stuart Kauffman
• Nealson KH, Conrad PG (December 1999). "Life: past, present and future". Philos. Trans. R.
Soc. Lond., B, Biol. Sci. 354 (1392): 1923–39. doi:10.1098/rstb.1999.0532. PMID 10670014.
PMC 1692713.
• Walker, Martin G. LIFE! Why We Exist...And What We Must Do to Survive Dog Ear
Publishing, 2006, ISBN 1-59858-243-7

[edit] External links

Wikimedia Commons has media related to: Tree of life
Wikiquote has a collection of quotations related to: Life

Look up life or living in Wiktionary, the free dictionary.

Wikispecies has information related to: The Taxonomy of Life

• Wikispecies - a free directory of life

• "The Adjacent Possible: A Talk with Stuart Kauffman"
• Stanford Encyclopedia of Philosophy entry
• Life under extreme conditions An in depth look at how life can form under the most extreme
conditions and circumstaces.

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Hierarchy of life · Origin of life · Life on Earth · Eukaryota (Plants/Flora,

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Biosphere > Ecosystem > Community (Biocoenosis) > Population > Organism > Organ system
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Population genetic
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Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir Vie (homonymie).

Cet article ne cite pas suffisamment ses sources (mars 2009).
Si vous connaissez le thème traité, merci d'indiquer les passages à sourcer avec
{{Référence souhaitée}} ou, mieux, incluez les références utiles en les liant aux notes de
bas de page. (Modifier l'article)
Les trois âges de la Vie et la Mort, peinture de Hans Baldung.
La vie est le nom donné :
1. à l'état et aux formes auto-organisées et homéostatiques de la matière (organismes vivants)
ayant une capacité de duplication et d'évolution. Cette définition est parfois étendue à l'ensemble des
êtres vivants dans la biosphère.
2. à un phénomène empirique particulièrement important pour les humains (qui sont eux-
mêmes vivants et pour qui les autres êtres vivants ont une place particulière), mais qui ne se laisse
pas facilement définir (cf. infra). Ce phénomène s'oppose à la notion de matière inerte ou inanimé qui
s'articule avec la notion de mort ;
3. à la durée qui sépare la naissance de la mort ;
4. au contenu en événements ou en actions de cette étendue temporelle, pour un humain ;
5. à l'approche harmonieuse des relations humaines (voir « question sociale »).
Une des marques de l'hominisation est l'existence de rites funéraires, et donc d'une conscience
d'une transition entre la vie et la mort. La vie est un concept primordial qui a donné lieu depuis des temps
immémoriaux à de nombreuses réflexions empiriques, philosophiques, scientifiques, etc. C'est également un
sujet de débat politique, qu'il s'agisse du traitement accordé aux êtres vivants par rapport aux humains et aux
choses inertes (cf. écologisme) ou des considérations sur le début et la fin de la vie humaine (cf. avortement,
euthanasie, « vie éternelle »), biophilie.
Ces réflexions concernent par exemple :
• la catégorie non-statique (par opposition à la matière inerte ou à l'état de mort) ;
• le concept d’évolution (passage de la matière inerte à la vie, développement et disparition
des formes vivantes, mort, création, etc.).
• la qualité de vie
Elles sont toujours liées aux notions d'esprit et d'intelligence. Elles débouchent également sur des
réflexions sur l'étendue temporelle et spatiale de la vie (y compris dans l'univers : « vie extraterrestre »).
Elles s'interrogent à la fois sur les conditions d'apparition de la vie (phénomène unique ou au contraire très
banal) et sur la possibilité d'une vie évoluée (par comparaison à l'humanité, implicitement considérée comme
l'achèvement de l'évolution de la vie terrestre) au sein de l'univers.
La biologie est l'étude scientifique de la vie. Elle s'appuie notamment sur la chimie organique mais
certains théoriciens n'excluent pas d'adopter des définitions pouvant inclure des formes mécaniques ou
électromécaniques, et même des formes créées par l'homme hors de tout processus reproductif naturel
(« vie artificielle » ou cellule artificielle).
• 1 Science
• 1.1 Domaine du vivant
• 1.2 Définitions
• 1.3 Origine de la vie
• 1.4 La vie comme propriété d’un organisme vivant
• 1.5 Caractéristiques biologiques d’un être vivant
• 1.5.1 Caractéristiques au niveau des
• 1.5.2 Caractéristiques au niveau des
structures et composés chimiques
• 1.5.3 Classification
• 1.5.4 Des organismes presque vivants
• 1.6 Autres définitions
• 2 Philosophie
• 2.1 Idéalisme et matérialisme
• 2.2 Une définition phénoménologique
• 3 Religion
• 3.1 Christianisme
• 4 Annexes
Science [modifier]
Domaine du vivant [modifier]
Le phénomène de la vie est potentiellement présent ailleurs dans l'Univers, mais n'est connu de
l'Homme que dans la biosphère terrestre. Elle est principalement concentrée dans les quelques centimètres
(ou premiers mètres) du sol, se produit surtout dans le premier mètre au-dessus du sol, ou les premiers
dizaines de mètres (mais se rencontre jusqu'à vers 10 km d'altitude) ainsi que dans les eaux douces et
marines. Mais on connait quelques formes de vies endolithiques : dans la roche, dans le pétrole ; et
extrémophiles : certaines faisant preuves d'une grande résistance (y compris au vide poussé, à la
radioactivité, à des pressions, pH ou températures extrêmes (chaud ou froid)).

Définitions [modifier]
Les expériences de Louis Pasteur ont réfuté la génération spontanée d'organismes vivants. Mais, il
n'existe pas encore de définition scientifique de la vie unanimement partagée. Toute définition doit tenir
compte de la notion de niveaux d'organisation structurels, d'émergence, d'homéostasie, d'entropie et de
métabolisme pour éviter de se retrouver dans une « zone grise ». Les définitions suivantes semblent limiter
ces zones grises :
• Selon la NASA, est vivant tout système délimité sur le plan spatial par une membrane semi-
perméable de sa propre fabrication et capable de s'auto-entretenir, ainsi que de se reproduire en
fabriquant ses propres constituants à partir d'énergie et/ou à partir d'éléments extérieurs.
• La vie est un état organisé et homéostatique de la matière.
• Mode d’organisation de la matière générant des formes diverses, de complexités variables,
en interaction et ayant comme propriété principale de se reproduire presque à l’identique en utilisant
les matériaux et l'énergie disponibles dans leur environnement, auquel elles peuvent s’adapter.
L'expression presque à l’identique réfère aux mutations qui apparaissent lors de la réplication de
l'organisme et qui peuvent conférer un avantage adaptatif à celui-ci.

Origine de la vie [modifier]

Article détaillé : origine de la vie.
L'origine de la vie et la relation entre ses lignées majeures font l'objet de recherches incessantes,
sans cesse bouleversées par de nouvelles découvertes scientifiques, en particulier la biologie moléculaire
pour ces dernières années.
On peut distinguer trois principaux groupes, les procaryotes, les eucaryotes et les archaeas. Deux
organites symbiotiques trouvés chez les eucaryotiques, à savoir la mitochondrie animale et le chloroplaste
végétal, sont considérés comme le résultat de l'endosymbiose d'une bactérie.

La vie comme propriété d’un organisme vivant [modifier]

L’organisme vivant est l’objet d’un processus de développement, la vie, qui le conduit en général par
étapes d’un état « embryonnaire » à l’adulte et à la mort, de manière individuelle ou coloniale, libre ou fixée,
tout ou partie de sa vie.
La graine, la spore, le spermatozoïde ou l’ovule sont aussi des formes du vivant, bien qu’ils n’aient
en eux-mêmes ni la forme ni les caractéristiques des êtres vivants qu’ils vont devenir. Il est ainsi difficile
d’isoler totalement la vie d’un individu de la lignée à laquelle il appartient et de la biosphère. Le vivant nait du
vivant : nous ne connaissons pas de vivant émergeant de l'inerte, ce qui rend difficile la reconstitution des
étapes prébiotiques.

Caractéristiques biologiques d’un être vivant [modifier]

Ou, comment peut-on affirmer qu’une entité est « vivante » ?

Caractéristiques au niveau des activités [modifier]

En biologie, une entité est traditionnellement considérée comme vivante si elle présente les activités
suivantes, au moins une fois durant son existence :
1. Développement ou croissance : l’entité grandit ou mûrit jusqu’au moment où elle devient
capable de se reproduire ;
2. Métabolisme : consommation, transformation et stockage d'énergie ou de masse; croissance
en absorbant de l’énergie ou des nutriments présents dans son environnement ou en réorganisant sa
masse, par production d’énergie, de travail et rejet de déchets ;
3. Motricité externe (locomotion) ou interne (circulation) ;
4. Reproduction : pouvoir créer de façon autonome d'autres entités similaires à soi-même.
5. Réponse à des stimuli : pouvoir détecter des propriétés de son environnement et d'agir de
façon adaptée.
Discussion sur ces critères :
• Ils ne sont pas tous satisfaits en même temps pour un individu particulier : il faut parfois
considérer la lignée ou l’espèce pour qu’ils coexistent (les hybrides stériles sont des êtres vivants) ;
• En isoler un ou deux peut conduire à des conclusions erronées : le feu (combustion)
assimilable à une digestion, car ce sont deux processus d’oxydation, ne transforme pas le feu en être
vivant ;
• Parfois, un critère manque : les virus ne grandissent pas et n'ont pas d'activité métabolique,
mais certains les considèrent comme vivants puisqu’ils peuvent contenir de l’ADN et être munis de
mécanismes (transcription d’ADN en ARN) provoquant leur reproduction dans les cellules hôtes ;
• D’autres fois encore, c’est une seule propriété qui est présente et qui se transmet à d’autres
entités, comme un mime de la fonction de reproduction (le prion est une protéine, conformée en
miroir par rapport à la protéine normale, qui transmet sa propriété pathogène aux autres protéines),

Caractéristiques au niveau des structures et composés chimiques [modifier]

D’où le besoin, éprouvé par les biologistes, de compléter ces caractéristiques pour réduire ces
ambiguïtés :
1. Les organismes vivants sont composés au moins d'une cellule, c’est-à-dire d’une membrane
fermée, séparant le milieu extérieur du milieu intérieur, qui contient le métabolisme et le matériel
2. Les organismes vivants contiennent des molécules telles que : des hydrates de carbone, des
lipides, des acides nucléiques et des protéines, toutes à base de carbone ; mais on peut y voir une
vision biaisée parce que carbocentrique de la vie. Des formes de vie pourraient en théorie être
fondées sur le silicium, mais celui-ci ne présente pas l’étonnante variété de formes et de propriétés
du carbone.
Selon la source d'énergie utilisée, on distingue les chimiotrophes, tirant leur énergie de molécules et
les Phototrophes, tirant leur énergie du Soleil.
Un organisme vivant est un ensemble organisé de matière qui tend à se maintenir à l'état
homéostatique par une utilisation concertée d'énergie.

Classification [modifier]
Articles connexes : Classification scientifique des espèces, Classification phylogénétique et
Classification classique.
Les organismes vivants sont classés par la phylogénétique. La première subdivision comprend trois
domaines :
• les archéobactéries ;
• les eubactéries ;
• les eucaryotes.

Des organismes presque vivants [modifier]

Il existe des entités proches des organismes vivants, qui ne sont toutefois pas considérés comme
tels, du fait d'une absence de métabolisme : les virus et les prions. Cependant, ces entités partagent avec les
organismes vivants la capacité de se répliquer, c'est-à-dire de susciter de la part de leur environnement la
production de copies d'elles-mêmes (formulation de David Deutsch) : ce sont des réplicateurs.

Autres définitions [modifier]

Pour Francisco Varela et Humberto Maturana, une entité est vivante si :
• elle peut se reproduire elle-même ;
• elle est basée sur l'eau ;
• elle produit des lipides et des protéines (?) ;
• son métabolisme est basé sur le carbone ;
• elle se réplique grâce à des acides nucléiques ;
• elle possède un système permettant de « lire » des protéines.
Cette définition a été largement utilisée par Lynn Margulis.
« Un système de rétrocontrôles négatifs inférieurs subordonnés à un rétrocontrôle positif supérieur »
(J. theor Biol. 2001)
Tom Kinch définit la vie comme un système autophage, hautement organisé, émergeant
naturellement des conditions ordinaires sur les corps planétaires et qui consiste en une population de
réplicateurs capables de muter.
Dans L'aventure du vivant, le biologiste Joël de Rosnay énumère trois propriétés fondamentales :
• L'autoconservation, qui est la capacité des organismes à se maintenir en vie par
l'assimilation, la nutrition, les réactions énergétiques de fermentation et de respiration ;
• L'autoreproduction, leur possibilité de propager la vie ;
• L'autorégulation : les fonctions de coordination, de synchronisation et de contrôle des
réactions d'ensemble.
Il faut ajouter à ces trois propriétés la capacité des êtres vivants à évoluer.

Philosophie [modifier]
La vie est un système ordonné capable de volonté. Ce système est dépendant de l'environnement
dans lequel il évolue.

Idéalisme et matérialisme [modifier]

Deux grands groupes de définitions sont discutés depuis les débuts de la philosophie : les
conceptions idéalistes qui s’appuient sur une séparation plus ou moins nette entre la matière et la vie (cf. la
définition phénoménologique, ci-après) et les conceptions matérialistes qui supposent la vie comme une des
manifestations émergentes de la matière.
Historiquement, on peut distinguer deux thèses, sans qu'il soit possible de déterminer si l'une est
antérieure à l'autre, d'autant qu'elles peuvent faire l'objet de synthèses variées (les deux thèses cohabitant à
des degrés divers au sein de théories plus sophistiquées). On les retrouve dans la pensée grecque antique.
Selon les thèses dites dualistes, la vie est conçue comme fondamentalement différente de la
matière : il y a du vivant (spirituel) et de l'inerte (matériel et énergie) comme il y a du fer et de l'eau. La seule
difficulté, c'est de « purifier » et « d'isoler » (au sens quasiment chimique) le vivant de l'inerte, séparation
d'autant plus difficile qu'elle est, par définition, inaccessible aux méthodes exclusivement matérielles. Ces
thèses font appel à des notions diverses : l’âme, le souffle vital, l’élan vital, etc. Cette séparation a donné lieu
à diverses théories, comme celle de la génération spontanée, encore vivaces au temps de Louis Pasteur.
Selon les thèses monistes, au contraire, la vie est une manifestation de la matière, une propriété
émergente qui apparaît spontanément dans certaines conditions. Il est alors possible de faire varier la
définition de la vie selon les conditions qu'on considère comme caractéristiques, ce qui introduit des marges
de faux débats (les contradicteurs croyant discuter sur le concept de vie alors que, en adoptant des critères
différents, ils s'interdisent a priori tout accord) même si en pratique seuls les objets en marge sont sujet à
discussion (les microbes, les virus, les prions, le feu, etc.). La pensée scientifique moderne relève de ce type
de thèse, en particulier suite aux expériences de Pasteur sur la stérilisation : tant qu'on n'a pas démontré la
nécessité de postuler une dualité, il convient de s'en tenir à l'hypothèse moniste. Même si les étapes de
l’apparition de la vie, ou de l'organisation des êtres vivants, restent à expliquer, les lois chimiques connues
sont pour l'instant suffisantes.
Les recherches sur les conditions matérielles originelles de notre planète, avec l’espoir de parvenir à
croiser ces informations avec celles existant sur d’autres planètes, nous donneront peut-être un jour un ou
des scénarios convaincants du passage de la matière inerte à la vie.

Une définition phénoménologique [modifier]

Article détaillé : phénoménologie de la vie.
Le philosophe Michel Henry définit la vie d'un point de vue phénoménologique comme ce qui
possède la faculté et le pouvoir « de se sentir et de s'éprouver soi-même en tout point de son être ». Pour lui,
la vie est essentiellement force subjective et affectivité, elle consiste en une pure expérience subjective de
soi qui oscille en permanence entre la souffrance et la joie. Une « force subjective » n’est pas une force
impersonnelle, aveugle et insensible comme le sont les forces objectives que l’on rencontre dans la nature,
mais une force vivante et sensible éprouvée de l’intérieur et résultant d’un désir subjectif et d’un effort
subjectif de la volonté pour le satisfaire. Il établit également une opposition radicale entre la chair vivante
douée de sensibilité et le corps matériel, qui est par principe insensible, dans son livre Incarnation, une
philosophie de la chair.

Religion [modifier]
Christianisme [modifier]
La religion insiste sur le caractère inaliénable de la vie en tant que fruit de la création divine. Le livre
de la Genèse contient le récit de la création.
Dans les dix commandements, il est écrit qu'il est interdit de tuer. Le décalogue est en quelque sorte
un code de vie pour les Israélites et, dans un certain sens, pour les Chrétiens également.
Dans le Nouveau Testament, Jésus dit Je suis la voie, la vérité et la vie. (Jn 14, 6). L'Esprit Saint est
appelé souffle de vie. La vie surnaturelle trouve sa source dans l'union hypostatique de Dieu.
Le magistère a adressé les encycliques : Evangelium vitae et Humanae Vitae, sur le droit à la vie et,
au respect fondamental qu'il lui est dû.

Annexes [modifier]
Sur les autres projets Wikimédia :
• Vie sur le Wiktionnaire (dictionnaire universel)
• Vie sur Wikisource (bibliothèque universelle)
• Vie sur Wikiquote (recueil de citations)

Articles connexes [modifier]

• Origines de la vie
• Histoire évolutive du vivant
• Mort - Fin de vie
• Vie après la mort
• Demi-vie
• Valeur de la vie
• Vérité Compréhension de la vie

• Portail de la médecine

• Portail de la philosophie
Ce document provient de « ».

Catégories : Vivant | Concept philosophique | [+]


Cell (biology)
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Drawing of the structure of cork as it appeared under the microscope to Robert Hooke from
Micrographia which is the origin of the word "cell" being used to describe the smallest unit of a living

Cells in culture, stained for keratin (red) and DNA (green)

The cell is the functional basic unit of life. It was discovered by Robert Hooke and is the functional
unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often
called the building block of life.[1] Some organisms, such as most bacteria, are unicellular (consist of a single
cell). Other organisms, such as humans, are multicellular. Humans have about 100 trillion or 1014 cells; a
typical cell size is 10 µm and a typical cell mass is 1 nanogram. The largest cells are about 135 µm in the
anterior horn in the spinal cord while granule cells in the cerebellum, the smallest, can be some 4 µm and the
longest cell can reach from the toe to the lower brain stem (Pseudounipolar cells).[2] The largest known cells
are unfertilised ostrich egg cells which weigh 3.3 pounds.[3][4]
In 1835, before the final cell theory was developed, Jan Evangelista Purkyně observed small
"granules" while looking at the plant tissue through a microscope. The cell theory, first developed in 1839 by
Matthias Jakob Schleiden and Theodor Schwann, states that all organisms are composed of one or more
cells, that all cells come from preexisting cells, that vital functions of an organism occur within cells, and that
all cells contain the hereditary information necessary for regulating cell functions and for transmitting
information to the next generation of cells.[5]
The word cell comes from the Latin cellula, meaning, a small room. The descriptive term for the
smallest living biological structure was coined by Robert Hooke in a book he published in 1665 when he
compared the cork cells he saw through his microscope to the small rooms monks lived in.[6]
• 1 Anatomy
• 1.1 Prokaryotic cells
• 1.2 Eukaryotic cells
• 2 Subcellular components
• 2.1 Membrane
• 2.2 Cytoskeleton
• 2.3 Genetic material
• 2.4 Organelles
• 3 Structures outside the cell wall
• 3.1 Capsule
• 3.2 Flagella
• 3.3 Fimbriae (pili)
• 4 Functions
• 4.1 Growth and metabolism
• 4.2 Creation
• 4.3 Protein synthesis
• 5 Movement or motility
• 6 Evolution
• 6.1 Origin of the first cell
[edit] Anatomy
There are two types of cells: eukaryotic and prokaryotic. Prokaryotic cells are usually independent,
while eukaryotic cells are often found in multicellular organisms.

[edit] Prokaryotic cells

Main article: Prokaryote
Diagram of a typical prokaryotic cell
The prokaryote cell is simpler, and therefore smaller, than a eukaryote cell, lacking a nucleus and
most of the other organelles of eukaryotes. There are two kinds of prokaryotes: bacteria and archaea; these
share a similar structure.
Nuclear material of prokaryotic cell consist of a single chromosome which is in direct contact with
cytoplasm. Here the undefined nuclear region in the cytoplasm is called nucleoid.
A prokaryotic cell has three architectural regions:
• On the outside, flagella and pili project from the cell's surface. These are structures (not
present in all prokaryotes) made of proteins that facilitate movement and communication between
• Enclosing the cell is the cell envelope – generally consisting of a cell wall covering a plasma
membrane though some bacteria also have a further covering layer called a capsule. The envelope
gives rigidity to the cell and separates the interior of the cell from its environment, serving as a
protective filter. Though most prokaryotes have a cell wall, there are exceptions such as Mycoplasma
(bacteria) and Thermoplasma (archaea). The cell wall consists of peptidoglycan in bacteria, and acts
as an additional barrier against exterior forces. It also prevents the cell from expanding and finally
bursting (cytolysis) from osmotic pressure against a hypotonic environment. Some eukaryote cells
(plant cells and fungi cells) also have a cell wall;
• Inside the cell is the cytoplasmic region that contains the cell genome (DNA) and ribosomes
and various sorts of inclusions. A prokaryotic chromosome is usually a circular molecule (an
exception is that of the bacterium Borrelia burgdorferi, which causes Lyme disease). Though not
forming a nucleus, the DNA is condensed in a nucleoid. Prokaryotes can carry extrachromosomal
DNA elements called plasmids, which are usually circular. Plasmids enable additional functions, such
as antibiotic resistance.

[edit] Eukaryotic cells

Main article: Eukaryote
Diagram of a typical animal (eukaryotic) cell, showing subcellular components.
(1) nucleolus
(2) nucleus
(3) ribosome
(4) vesicle
(5) rough endoplasmic reticulum (ER)
(6) Golgi apparatus
(7) Cytoskeleton
(8) smooth endoplasmic reticulum
(9) mitochondria
(10) vacuole
(11) cytoplasm
(12) lysosome
(13) centrioles within centrosome
Eukaryotic cells are about 15 times wider than a typical prokaryote and can be as much as 1000
times greater in volume. The major difference between prokaryotes and eukaryotes is that eukaryotic cells
contain membrane-bound compartments in which specific metabolic activities take place. Most important
among these is a cell nucleus, a membrane-delineated compartment that houses the eukaryotic cell's DNA.
This nucleus gives the eukaryote its name, which means "true nucleus." Other differences include:
• The plasma membrane resembles that of prokaryotes in function, with minor differences in
the setup. Cell walls may or may not be present.
• The eukaryotic DNA is organized in one or more linear molecules, called chromosomes,
which are associated with histone proteins. All chromosomal DNA is stored in the cell nucleus,
separated from the cytoplasm by a membrane. Some eukaryotic organelles such as mitochondria
also contain some DNA.
• Many eukaryotic cells are ciliated with primary cilia. Primary cilia play important roles in
chemosensation, mechanosensation, and thermosensation. Cilia may thus be "viewed as sensory
cellular antennae that coordinate a large number of cellular signaling pathways, sometimes coupling
the signaling to ciliary motility or alternatively to cell division and differentiation."[7]
• Eukaryotes can move using motile cilia or flagella. The flagella are more complex than those
of prokaryotes.
Table 1: Comparison of features of prokaryotic and eukaryotic cells
Prokaryotes Eukaryotes

Typical organisms bacteria, archaea protists, fungi, plants, animals

~ 10–100 µm (sperm cells, apart from the

Typical size ~ 1–10 µm
tail, are smaller)

nucleoid region; no
Type of nucleus real nucleus with double membrane
real nucleus

DNA circular (usually) linear molecules (chromosomes) with

histone proteins

RNA-/protein- coupled in RNA-synthesis inside the nucleus

synthesis cytoplasm protein synthesis in cytoplasm

Ribosomes 50S+30S 60S+40S

Cytoplasmatic highly structured by endomembranes and

very few structures
structure a cytoskeleton

flagella made of flagella and cilia containing microtubules;

Cell movement
flagellin lamellipodia and filopodia containing actin

one to several thousand (though some

Mitochondria none
lack mitochondria)

Chloroplasts none in algae and plants

single cells, colonies, higher multicellular

Organization usually single cells
organisms with specialized cells

Binary fission Mitosis (fission or budding)

Cell division
(simple division) Meiosis
Table 2: Comparison of structures between animal and plant cells
Typical animal cell Typical plant cell

• Nucleus
• Nucleus
• Nucleolus (within
• Nucleolus (within
• Rough endoplasmic
• Rough ER
reticulum (ER)
• Smooth ER
• Smooth ER
• Ribosomes
• Ribosomes
• Cytoskeleton
Organelles • Cytoskeleton
• Golgi apparatus
• Golgi apparatus
• Cytoplasm
• Cytoplasm
• Mitochondria
• Mitochondria
• Vesicles
• Plastids and its derivatives
• Lysosomes
• Vacuole(s)
• Centrosome
• Cell wall
• Centrioles
[edit] Subcellular components

The cells of eukaryotes (left) and prokaryotes (right)

All cells, whether prokaryotic or eukaryotic, have a membrane that envelops the cell, separates its
interior from its environment, regulates what moves in and out (selectively permeable), and maintains the
electric potential of the cell. Inside the membrane, a salty cytoplasm takes up most of the cell volume. All
cells possess DNA, the hereditary material of genes, and RNA, containing the information necessary to build
various proteins such as enzymes, the cell's primary machinery. There are also other kinds of biomolecules
in cells. This article will list these primary components of the cell, then briefly describe their function.

[edit] Membrane
Main article: Cell membrane
The cytoplasm of a cell is surrounded by a cell membrane or plasma membrane. The plasma
membrane in plants and prokaryotes is usually covered by a cell wall. This membrane serves to separate
and protect a cell from its surrounding environment and is made mostly from a double layer of lipids
(hydrophobic fat-like molecules) and hydrophilic phosphorus molecules. Hence, the layer is called a
phospholipid bilayer. It may also be called a fluid mosaic membrane. Embedded within this membrane is a
variety of protein molecules that act as channels and pumps that move different molecules into and out of the
cell. The membrane is said to be 'semi-permeable', in that it can either let a substance (molecule or ion) pass
through freely, pass through to a limited extent or not pass through at all. Cell surface membranes also
contain receptor proteins that allow cells to detect external signaling molecules such as hormones.

[edit] Cytoskeleton
Main article: Cytoskeleton
Bovine Pulmonary Artery Endothelial cell: nuclei stained blue, mitochondria stained red, and F-actin,
an important component in microfilaments, stained green. Cell imaged on a fluorescent microscope.
The cytoskeleton acts to organize and maintain the cell's shape; anchors organelles in place; helps
during endocytosis, the uptake of external materials by a cell, and cytokinesis, the separation of daughter
cells after cell division; and moves parts of the cell in processes of growth and mobility. The eukaryotic
cytoskeleton is composed of microfilaments, intermediate filaments and microtubules. There is a great
number of proteins associated with them, each controlling a cell's structure by directing, bundling, and
aligning filaments. The prokaryotic cytoskeleton is less well-studied but is involved in the maintenance of cell
shape, polarity and cytokinesis.[8]

[edit] Genetic material

Two different kinds of genetic material exist: deoxyribonucleic acid (DNA) and ribonucleic acid
(RNA). Most organisms use DNA for their long-term information storage, but some viruses (e.g., retroviruses)
have RNA as their genetic material. The biological information contained in an organism is encoded in its
DNA or RNA sequence. RNA is also used for information transport (e.g., mRNA) and enzymatic functions
(e.g., ribosomal RNA) in organisms that use DNA for the genetic code itself. Transfer RNA (tRNA) molecules
are used to add amino acids during protein translation.
Prokaryotic genetic material is organized in a simple circular DNA molecule (the bacterial
chromosome) in the nucleoid region of the cytoplasm. Eukaryotic genetic material is divided into different,
linear molecules called chromosomes inside a discrete nucleus, usually with additional genetic material in
some organelles like mitochondria and chloroplasts (see endosymbiotic theory).
A human cell has genetic material contained in the cell nucleus (the nuclear genome) and in the
mitochondria (the mitochondrial genome). In humans the nuclear genome is divided into 23 pairs of linear
DNA molecules called chromosomes. The mitochondrial genome is a circular DNA molecule distinct from the
nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes, it codes for
13 proteins involved in mitochondrial energy production and specific tRNAs.
Foreign genetic material (most commonly DNA) can also be artificially introduced into the cell by a
process called transfection. This can be transient, if the DNA is not inserted into the cell's genome, or stable,
if it is. Certain viruses also insert their genetic material into the genome.

[edit] Organelles
Main article: Organelle
The human body contains many different organs, such as the heart, lung, and kidney, with each
organ performing a different function. Cells also have a set of "little organs," called organelles, that are
adapted and/or specialized for carrying out one or more vital functions. Both eukaryotic and prokaryotic cells
have organelles but organelles in eukaryotes are generally more complex and may be membrane bound.
There are several types of organelles in a cell. Some (such as the nucleus and golgi apparatus) are
typically solitary, while others (such as mitochondria, peroxisomes and lysosomes) can be numerous
(hundreds to thousands). The cytosol is the gelatinous fluid that fills the cell and surrounds the organelles.
Cell nucleus – eukaryotes only - a cell's information
The cell nucleus is the most conspicuous
organelle found in a eukaryotic cell. It houses the cell's
chromosomes, and is the place where almost all DNA
replication and RNA synthesis (transcription) occur.
The nucleus is spherical and separated from the
cytoplasm by a double membrane called the nuclear
envelope. The nuclear envelope isolates and protects
a cell's DNA from various molecules that could
accidentally damage its structure or interfere with its
processing. During processing, DNA is transcribed, or
copied into a special RNA, called messenger RNA
(mRNA). This mRNA is then transported out of the
nucleus, where it is translated into a specific protein
molecule. The nucleolus is a specialized region within
the nucleus where ribosome subunits are assembled.
Diagram of a cell nucleus
In prokaryotes, DNA processing takes place in the

Mitochondria are self-replicating organelles

that occur in various numbers, shapes, and sizes in
the cytoplasm of all eukaryotic cells. Mitochondria play
a critical role in generating energy in the eukaryotic
cell. Mitochondria generate the cell's energy by
oxidative phosphorylation, using oxygen to release
energy stored in cellular nutrients (typically pertaining
to glucose) to generate ATP. Mitochondria multiply by
splitting in two. Respiration occurs in the cell
Organelles that are modified chloroplasts are
broadly called plastids, and are involved in energy
storage through photosynthesis, which uses solar
energy to generate carbohydrates and oxygen from
carbon dioxide and water.[citation needed]
Mitochondria and chloroplasts each contain
their own genome, which is separate and distinct from
the nuclear genome of a cell. Both organelles contain
this DNA in circular plasmids, much like prokaryotic
cells, strongly supporting the evolutionary theory of
endosymbiosis; since these organelles contain their
own genomes and have other similarities to
prokaryotes, they are thought to have developed
through a symbiotic relationship after being engulfed
by a primitive cell.[citation needed]
Endoplasmic reticulum – eukaryotes only
The endoplasmic reticulum (ER) is the
transport network for molecules targeted for certain
modifications and specific destinations, as compared
to molecules that will float freely in the cytoplasm. The
ER has two forms: the rough ER, which has ribosomes
on its surface and secretes proteins into the
cytoplasm, and the smooth ER, which lacks them.
Smooth ER plays a role in calcium sequestration and
Golgi apparatus – eukaryotes only
The primary function of the Golgi apparatus is
to process and package the macromolecules such as
proteins and lipids that are synthesized by the cell. It is
particularly important in the processing of proteins for
secretion. The Golgi apparatus forms a part of the
endomembrane system of eukaryotic cells. Vesicles
that enter the Golgi apparatus are processed in a cis
to trans direction, meaning they coalesce on the cis
side of the apparatus and after processing pinch off on
the opposite (trans) side to form a new vesicle in the
animal cell.[citation needed]

Diagram of an endomembrane system

The ribosome is a large complex of RNA and
protein molecules. They each consist of two subunits,
and act as an assembly line where RNA from the
nucleus is used to synthesise proteins from amino
acids. Ribosomes can be found either floating freely or
bound to a membrane (the rough endoplasmatic
reticulum in eukaryotes, or the cell membrane in

Lysosomes and Peroxisomes – eukaryotes only

Lysosomes contain digestive enzymes (acid hydrolases). They digest excess or worn-out
organelles, food particles, and engulfed viruses or bacteria. Peroxisomes have enzymes that rid the
cell of toxic peroxides. The cell could not house these destructive enzymes if they were not contained
in a membrane-bound system. These organelles are often called a "suicide bag" because of their
ability to detonate and destroy the cell.[citation needed]

Centrosome – the cytoskeleton organiser

The centrosome produces the microtubules of a cell – a key component of the cytoskeleton. It
directs the transport through the ER and the Golgi apparatus. Centrosomes are composed of two
centrioles, which separate during cell division and help in the formation of the mitotic spindle. A single
centrosome is present in the animal cells. They are also found in some fungi and algae cells.[ citation
Vacuoles store food and waste. Some vacuoles store extra water. They are often described
as liquid filled space and are surrounded by a membrane. Some cells, most notably Amoeba, have
contractile vacuoles, which can pump water out of the cell if there is too much water. The vacuoles of
eukaryotic cells are usually larger in those of plants than animals.

[edit] Structures outside the cell wall

[edit] Capsule
A gelatinous capsule is present in some bacteria outside the cell wall. The capsule may be
polysaccharide as in pneumococci, meningococci or polypeptide as Bacillus anthracis or hyaluronic acid as
in streptococci.[citation needed] Capsules are not marked by ordinary stain and can be detected by special
stain. The capsule is antigenic. The capsule has antiphagocytic function so it determines the virulence of
many bacteria. It also plays a role in attachment of the organism to mucous membranes.[ citation needed]

[edit] Flagella
Flagella are the organelles of cellular mobility. They arise from cytoplasm and extrude through the
cell wall. They are long and thick thread-like appendages, protein in nature. Are most commonly found in
bacteria cells but are found in animal cells as well.

[edit] Fimbriae (pili)

They are short and thin hair like filaments, formed of protein called pilin (antigenic). Fimbriae are
responsible for attachment of bacteria to specific receptors of human cell (adherence). There are special
types of pili called (sex pili) involved in conjunction.[ citation needed]
[edit] Functions
[edit] Growth and metabolism
Main articles: Cell growth and Metabolism
Between successive cell divisions, cells grow through the functioning of cellular metabolism. Cell
metabolism is the process by which individual cells process nutrient molecules. Metabolism has two distinct
divisions: catabolism, in which the cell breaks down complex molecules to produce energy and reducing
power, and anabolism, in which the cell uses energy and reducing power to construct complex molecules
and perform other biological functions. Complex sugars consumed by the organism can be broken down into
a less chemically complex sugar molecule called glucose. Once inside the cell, glucose is broken down to
make adenosine triphosphate (ATP), a form of energy, through two different pathways.
The first pathway, glycolysis, requires no oxygen and is referred to as anaerobic metabolism. Each
reaction is designed to produce some hydrogen ions that can then be used to make energy packets (ATP). In
prokaryotes, glycolysis is the only method used for converting energy.
The second pathway, called the Krebs cycle, or citric acid cycle, occurs inside the mitochondria and
can generate enough ATP to run all the cell functions.
An overview of protein synthesis.
Within the nucleus of the cell (light blue), genes (DNA, dark blue) are transcribed into RNA. This RNA is then
subject to post-transcriptional modification and control, resulting in a mature mRNA (red) that is then
transported out of the nucleus and into the cytoplasm (peach), where it undergoes translation into a protein.
mRNA is translated by ribosomes (purple) that match the three-base codons of the mRNA to the three-base
anti-codons of the appropriate tRNA. Newly synthesized proteins (black) are often further modified, such as
by binding to an effector molecule (orange), to become fully active.

[edit] Creation
Main article: Cell division
Cell division involves a single cell (called a mother cell) dividing into two daughter cells. This leads to
growth in multicellular organisms (the growth of tissue) and to procreation (vegetative reproduction) in
unicellular organisms.
Prokaryotic cells divide by binary fission. Eukaryotic cells usually undergo a process of nuclear
division, called mitosis, followed by division of the cell, called cytokinesis. A diploid cell may also undergo
meiosis to produce haploid cells, usually four. Haploid cells serve as gametes in multicellular organisms,
fusing to form new diploid cells.
DNA replication, or the process of duplicating a cell's genome, is required every time a cell divides.
Replication, like all cellular activities, requires specialized proteins for carrying out the job.
[edit] Protein synthesis
Main article: Protein biosynthesis
Cells are capable of synthesizing new proteins, which are essential for the modulation and
maintenance of cellular activities. This process involves the formation of new protein molecules from amino
acid building blocks based on information encoded in DNA/RNA. Protein synthesis generally consists of two
major steps: transcription and translation.
Transcription is the process where genetic information in DNA is used to produce a complementary
RNA strand. This RNA strand is then processed to give messenger RNA (mRNA), which is free to migrate
through the cell. mRNA molecules bind to protein-RNA complexes called ribosomes located in the cytosol,
where they are translated into polypeptide sequences. The ribosome mediates the formation of a polypeptide
sequence based on the mRNA sequence. The mRNA sequence directly relates to the polypeptide sequence
by binding to transfer RNA (tRNA) adapter molecules in binding pockets within the ribosome. The new
polypeptide then folds into a functional three-dimensional protein molecule.

[edit] Movement or motility

Cells can move during many processes: such as wound healing, the immune response and cancer
metastasis. For wound healing to occur, white blood cells and cells that ingest bacteria move to the wound
site to kill the microorganisms that cause infection.
At the same time fibroblasts (connective tissue cells) move there to remodel damaged structures. In the case
of tumor development, cells from a primary tumor move away and spread to other parts of the body. Cell
motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins.[10] The
process is divided into three steps – protrusion of the leading edge of the cell, adhesion of the leading edge
and de-adhesion at the cell body and rear, and cytoskeletal contraction to pull the cell forward. Each step is
driven by physical forces generated by unique segments of the cytoskeleton.[11][12]

[edit] Evolution
Main article: Evolutionary history of life
The origin of cells has to do with the origin of life, which began the history of life on Earth.

[edit] Origin of the first cell

Further information: Abiogenesis
There are several theories about the origin of small molecules that could lead to life in an early Earth.
One is that they came from meteorites (see Murchison meteorite). Another is that they were created at deep-
sea vents. A third is that they were synthesized by lightning in a reducing atmosphere ( see Miller–Urey
experiment); although it is not clear if Earth had such an atmosphere. There are essentially no experimental
data defining what the first self-replicating forms were. RNA is generally assumed to be the earliest self-
replicating molecule, as it is capable of both storing genetic information and catalyzing chemical reactions
(see RNA world hypothesis). But some other entity with the potential to self-replicate could have preceded
RNA, like clay or peptide nucleic acid.[13]
Cells emerged at least 4.0–4.3 billion years ago. The current belief is that these cells were
heterotrophs. An important characteristic of cells is the cell membrane, composed of a bilayer of lipids. The
early cell membranes were probably more simple and permeable than modern ones, with only a single fatty
acid chain per lipid. Lipids are known to spontaneously form bilayered vesicles in water, and could have
preceded RNA. But the first cell membranes could also have been produced by catalytic RNA, or even have
required structural proteins before they could form.[14]

[edit] Origin of eukaryotic cells

The eukaryotic cell seems to have evolved from a symbiotic community of prokaryotic cells. DNA-
bearing organelles like the mitochondria and the chloroplasts are almost certainly what remains of ancient
symbiotic oxygen-breathing proteobacteria and cyanobacteria, respectively, where the rest of the cell seems
to be derived from an ancestral archaean prokaryote cell – a theory termed the endosymbiotic theory.
There is still considerable debate about whether organelles like the hydrogenosome predated the
origin of mitochondria, or viceversa: see the hydrogen hypothesis for the origin of eukaryotic cells.
Sex, as the stereotyped choreography of meiosis and syngamy that persists in nearly all extant
eukaryotes, may have played a role in the transition from prokaryotes to eukaryotes. An 'origin of sex as
vaccination' theory suggests that the eukaryote genome accreted from prokaryan parasite genomes in
numerous rounds of lateral gene transfer. Sex-as-syngamy (fusion sex) arose when infected hosts began
swapping nuclearized genomes containing co-evolved, vertically transmitted symbionts that conveyed
protection against horizontal infection by more virulent symbionts.[15]

[edit] History
• 1632–1723: Antonie van Leeuwenhoek teaches himself to grind lenses, builds a microscope
and draws protozoa, such as Vorticella from rain water, and bacteria from his own mouth.
• 1665: Robert Hooke discovers cells in cork, then in living plant tissue using an early
• 1839: Theodor Schwann and Matthias Jakob Schleiden elucidate the principle that plants
and animals are made of cells, concluding that cells are a common unit of structure and
development, and thus founding the cell theory.
• The belief that life forms can occur spontaneously (generatio spontanea) is contradicted by
Louis Pasteur (1822–1895) (although Francesco Redi had performed an experiment in 1668 that
suggested the same conclusion).
• 1855: Rudolf Virchow states that cells always emerge from cell divisions (omnis cellula ex
• 1931: Ernst Ruska builds first transmission electron microscope (TEM) at the University of
Berlin. By 1935, he has built an EM with twice the resolution of a light microscope, revealing
previously unresolvable organelles.
• 1953: Watson and Crick made their first announcement on the double-helix structure for DNA
on February 28.
• 1981: Lynn Margulis published Symbiosis in Cell Evolution detailing the endosymbiotic

[edit] See also

Wikimedia Commons has media related to: Cell biology

Main article: Topic outline of cell biology

• Cell biology
• Cell culture
• Cell type
• Cellular component
• Cytorrhysis
• Cytotoxicity
• Plasmolysis
• Stem cell
• Syncytium

[edit] References
1. ^ Cell Movements and the Shaping of the Vertebrate Body in Chapter 21 of Molecular
Biology of the Cell fourth edition, edited by Bruce Alberts (2002) published by Garland Science.
The Alberts text discusses how the "cellular building blocks" move to shape developing embryos. It is
also common to describe small molecules such as amino acids as "molecular building blocks".
2. ^ Integrative Biology 131 - Lecture 03: Skeletal System at YouTube first 12 minutes of the
lecture covers cells (by Marian Diamond).
3. ^ Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life.
Boston, Massachusetts: Pearson Prentice Hall. ISBN 0-13-250882-6.
4. ^ Mitzi Perdue. "Facts about Birds and Eggs". Retrieved
5. ^ Maton, Anthea; Hopkins, Jean Johnson, Susan LaHart, David Quon Warner, Maryanna
Wright, Jill D (1997). Cells Building Blocks of Life. New Jersey: Prentice Hall. ISBN 0-13-423476-6.
6. ^ a b "... I could exceedingly plainly perceive it to be all perforated and porous, much like a
Honey-comb, but that the pores of it were not regular [..] these pores, or cells, [..] were indeed the
first microscopical pores I ever saw, and perhaps, that were ever seen, for I had not met with any
Writer or Person, that had made any mention of them before this. . . " – Hooke describing his
observations on a thin slice of cork. Robert Hooke
7. ^ Satir, P; Christensen, ST; Søren T. Christensen (2008-03-26). "Structure and function of
mammalian cilia". Histochemistry and Cell Biology (Springer Berlin / Heidelberg) 129 (6): 687–693.
doi:10.1007/s00418-008-0416-9. 1432-119X. PMID 18365235. PMC 2386530. Retrieved 2009-09-12.
8. ^ Michie K, Löwe J (2006). "Dynamic filaments of the bacterial cytoskeleton". Annu Rev
Biochem 75: 467–92. doi:10.1146/annurev.biochem.75.103004.142452. PMID 16756499.
9. ^ Ménétret JF, Schaletzky J, Clemons WM, et al., CW; Akey (December 2007). "Ribosome
binding of a single copy of the SecY complex: implications for protein translocation". Mol. Cell 28 (6):
1083–92. doi:10.1016/j.molcel.2007.10.034. PMID 18158904.
10.^ Revathi Ananthakrishnan1 *, Allen Ehrlicher2 ✉. "The Forces Behind Cell Movement". Retrieved 2009-04-17.
11.^ Alberts B, Johnson A, Lewis J. et al. Molecular Biology of the Cell, 4e. Garland Science.
12.^ Ananthakrishnan R, Ehrlicher A. The Forces Behind Cell Movement. Int J Biol Sci 2007;
13.^ Orgel LE (1998). "The origin of life--a review of facts and speculations". Trends Biochem
Sci 23 (12): 491–5. doi:10.1016/S0968-0004(98)01300-0. PMID 9868373.
14.^ Griffiths G (December 2007). "Cell evolution and the problem of membrane topology".
Nature reviews. Molecular cell biology 8 (12): 1018–24. doi:10.1038/nrm2287. PMID 17971839.
15.^ Sterrer W (2002). "On the origin of sex as vaccination". Journal of Theoretical Biology 216:
387–396. doi:10.1006/jtbi.2002.3008. PMID 12151256.
• This article incorporates public domain material from the NCBI document "Science

[edit] External links

• Inside the Cell
• Virtual Cell's Educational Animations
• The Inner Life of A Cell, a flash video showing what happens inside of a cell
• The Virtual Cell
• Cells Alive!
• Journal of Cell Biology
• The Biology Project > Cell Biology
• Centre of the Cell online
• The Image & Video Library of The American Society for Cell Biology , a collection of peer-
reviewed still images, video clips and digital books that illustrate the structure, function and biology of
the cell.
[edit] Textbooks
• Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular Biology of the
Cell (4th ed.). Garland. ISBN 0815332181.
• Lodish H, Berk A, Matsudaira P, Kaiser CA, Krieger M, Scott MP, Zipurksy SL, Darnell J
(2004). Molecular Cell Biology (5th ed.). WH Freeman: New York, NY. ISBN 978-0716743668.
• Cooper GM (2000). The cell: a molecular approach (2nd ed.). Washington, D.C: ASM Press.
ISBN 0-87893-102-3.

Structures of the cell / organelles

Cell membrane · Nucleus (and Nucleolus) · Endoplasmic reticulum ·

Golgi apparatus · Parenthesome · Autophagosome
system Vesicles (Exosome · Lysosome · Endosome · Phagosome · Vacuole)
Cytoplasmic granules: Melanosome · Microbody (Glyoxysome,
Peroxisome) · Weibel-Palade body

Endosymbionts Mitochondrion · Plastids (Chloroplast · Chromoplast · Leucoplast)

Cytoskeleton Microfilaments · Intermediate filaments · Microtubules · Prokaryotic

MTOCs Centrosome/Centriole · Basal body · Spindle pole body

Undulipodium Cilium/Flagellum · Axoneme · Radial spoke

Other external Cell wall · Acrosome

Other Cytoplasm · Ribosome · Vault · Proteasome

B strc: perx, skel, epit, ctrs, cili, mito, nucl (chro)

Hierarchy of life

Biosphere > Ecosystem > Community (Biocoenosis) > Population > Organism > Organ system
> Organ > Tissue > Cell > Organelle > Molecule (Macromolecule · Biomolecule) > Atom
Retrieved from ""

Categories: Cell biology

• This page was last modified on 3 December 2010 at 07:06.


Cellule (biologie)
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir cellule.

Cellules épithéliales en culture. L'ADN est coloré en vert, les filaments de kératine en rouge.
Une petite section d'une membrane cellulaire. Cette membrane de cellule moderne est bien plus
sophistiquée que la simple phospholipide à deux couches originelle (les petites sphères à deux queues).
Protéines et glucides ont plusieurs fonctions de régulation du passage de matériau à travers la membrane et
de réaction à l'environnement.
La cellule (du latin cellula petite chambre) est l'unité structurale, fonctionnelle et reproductrice
constituant tout ou partie d'un être vivant (à l'exception des virus). Chaque cellule est une entité vivante qui,
dans le cas d'organismes multicellulaires, fonctionne de manière autonome, mais coordonnée avec les
autres. Les cellules de même type sont réunies en tissus, eux-mêmes réunis en organes.
La théorie cellulaire implique l'unité de tout le vivant : tous les êtres vivants sont composés de
cellules dont la structure fondamentale est commune ainsi que l'homéostasie du milieu intérieur, milieu de
composition physico-chimique régulé et propice au développement des cellules de l'espèce considérée.
• 1 Histoire du concept
• 1.1 Théorie cellulaire
• 2 La notion de cellule
• 2.1 La cellule, machine à produire de l'ordre
• 2.1.1 Un espace clos effectuant des
échanges avec l'extérieur
• 2.1.2 La compartimentation : mise en place
de microenvironnements aux propriétés spécifiques
• 2.1.3 La cellule un flux organisé de matière
et d'énergie
• 2.2 La cellule vectrice de gènes
• 2.2.1 La transmission des gènes et cycle
• 2.2.2 La reproduction sexuée et cycle du
• 2.2.3 La mort cellulaire : la cellule au
service de l'organisme et des gènes
• 2.3 Interdépendance cellulaire : de la cellule à
• 2.3.1 Les êtres unicellulaires : la cellule
Histoire du concept [modifier]

Dessin de « cellules » observées dans des coupes d'écorce d'arbre par Robert Hooke en 1665.
• 1665 : Robert Hooke découvre des cellules mortes dans du liège, ces cellules lui font penser
aux cellules d'un monastère, d'où le nom. Puis il observe des cellules dans des plantes vivantes, en
utilisant les premiers microscopes.
• 1839 : Theodor Schwann découvre que les plantes et les animaux sont tous faits de cellules,
concluant que la cellule est l'unité commune de structure et de développement, ce qui fonda la
théorie cellulaire. Il donna son nom aux cellules de Schwann.
• La croyance selon laquelle des formes de vie peuvent apparaître spontanément ( génération
spontanée) est réfutée par Louis Pasteur (1822-1895).
• 1858 : Rudolf Virchow affirma que les cellules naissent du résultat de la division cellulaire
(« omnis cellula ex cellula »), ce qui repose en termes cellulaires la question de l'œuf et de la poule.
C'est précisément cette partie qui est attaquée par les tenants du créationnisme ou de son dernier
avatar, le dessein intelligent.

Théorie cellulaire [modifier]

Article détaillé : Théorie cellulaire.
1. La cellule est l'unité constitutive des organismes vivants. Elle en est aussi l'unité
2. L'organisme dépend de l'activité des cellules isolées ou groupées en tissus pour assurer les
différentes fonctions.
3. Les activités biochimiques des cellules sont coordonnées et déterminées par certaines
structures présentes à l'intérieur des cellules.
4. La multiplication des cellules permet le maintien des organismes et leur multiplication.
5. Cette théorie est formulée en 1838 par Schleiden et Schwann : la cellule est unité de vie
(tout ce qui est vivant est cellulaire). Cette théorie évoque également la présence d'organites à
l'intérieur de ces mêmes cellules.

La notion de cellule [modifier]

Ici on se demande avant tout quelles sont les caractéristiques communes aux cellules, malgré leur

La cellule, machine à produire de l'ordre [modifier]

La cellule représente un état hautement organisé de la matière : maintenir cet ordre tout en étant
soumis aux principes de la thermodynamique nécessite la mise en place de structures permettant d'utiliser
l'énergie, la matière extérieure (on crée de l'ordre au niveau de la cellule mais, globalement, l'entropie
augmente) ; la cellule est donc un système thermodynamiquement ouvert.

Un espace clos effectuant des échanges avec l'extérieur [modifier]

La cellule constitue une unité spatiale, délimitée par une membrane. Celle-ci, loin d'être une limite
hermétique, constitue une surface d'échanges permettant la mise en place de flux.
Les membranes plasmiques, malgré leur diversité possèdent, sauf exceptions (certaines archées
thermophiles possédant une seule couche de lipides), une structure identique :
• une bicouche phospholipidique composée de lipides amphiphiles, qui constitue un filtre de
base permettant le passage des substances hydrophobes, freinant celui des hydrophiles.
• des protéines transmembranaires et périphériques aux rôles divers (transferts, transport,
transduction de signaux...)
La membrane agit non seulement comme un filtre, c'est-à-dire en laissant passer certaines
molécules selon la différence de concentration (appelée à tort gradient de concentration) mais aussi en
utilisant de l'énergie (osmotique, chimique...) pour favoriser les flux endergoniques. Elle permet aussi le
passage de la lumière, de la chaleur... En tant que surface de contact avec l'extérieur, elle assure aussi la
transmission d'informations nécessaires à la réactivité de la cellule aux changements de l'environnement et à
la coordination avec d'autres cellules.
La membrane plasmique crée donc un espace clos en constant échange avec l'environnement

La compartimentation : mise en place de microenvironnements aux propriétés

spécifiques [modifier]
La présence d'une membrane biologique entourant un espace, que ce soit le cytoplasme ou la
lumière d'un organite, va permettre, en contrôlant les échanges des macromolécules, des ions (et de toute
autre molécule) l'établissement de conditions favorisant certaines réactions par rapport à d'autres : en variant
les différents facteurs physico-chimiques (pH, concentration en ions...), la nature des enzymes et des
produits, leur nombre...
Cet environnement permet ainsi la biosynthèse et la dégradation de molécules organiques, et ainsi
le maintien d'une structure hautement organisée par un recyclage constant des molécules qui le forment.
Cette compartimentation se trouve particulièrement poussée dans le cas des eucaryotes : elle
permet la spécialisation fonctionnelle des différents organites (la composition de leur lumière étant différente,
ils sont le siège de réactions différentes : on va ainsi pouvoir favoriser la production de tel produit dans un
compartiment, sa destruction dans un autre).

La cellule un flux organisé de matière et d'énergie [modifier]

Cette structure de base (une membrane organisant les échanges entourant un compartiment, lieu de
réactions chimiques spécifiques) va permettre la mise en place et le maintien de flux de matière, d'énergie,
d'information... traversant la cellule. Il y a donc une réelle organisation des échanges cellule-extérieur, qui va
permettre au « système cellule » (au sens thermodynamique) de maintenir sa structure hautement
Finalement, la cellule ne constitue pas une structure stable et immuable mais plutôt une entité
dynamique nécessitant un apport constant de matière et d'énergie pour permettre son fonctionnement et le
maintien de sa structure : seule son organisation persiste, ses constituants étant en perpétuel
Le potentiel de repos de la cellule détermine la différence de potentiel (-70 mV
environ[réf. nécessaire]), avec l'intérieur de la cellule négatif.

La cellule vectrice de gènes [modifier]

Cette structure résulte de l'expression d'un programme génétique complexe (permettant notamment
la synthèse d'enzymes dont on vient de voir l'importance). Celui-ci doit être transmis, en même temps que la
structure de base, au cours des divisions cellulaires. La cellule peut donc être considérée non seulement
comme l'unité structurelle du vivant mais aussi comme un vecteur de gènes assurant leur transmission au fil
des générations.

La transmission des gènes et cycle cellulaire [modifier]

La division cellulaire aboutissant, à partir d'une cellule mère, à deux cellules filles contenant le même
génome (aux erreurs de copie près) nécessite la succession de phases de synthèse protéique permettant le
renouvellement et la croissance cellulaire, de synthèse d'ADN et enfin de partition plus ou moins équitable
de la cellule.
La synthèse protéique résulte de l'expression du matériel génétique, elle se déroule en plusieurs
étapes : transcription de l'ADN en ARN, traduction de l'ARN en une chaîne polypeptidique, repliement de
celle-ci (chez les eucaryotes s'insèrent des phases de maturation où l'on coupe et modifie la séquence
synthétisée). La copie du génome est réalisée par toute une machinerie protéique permettant à l'ADN
polymérase d'accéder à la séquence et de la copier, selon le principe d'appariement des bases.
La partition de la cellule se fait par des mécanismes différents chez les procaryotes et les eucaryotes
(nommée alors mitose) : celle-ci consiste en la partition et la transmission du génome intégral de la cellule

La reproduction sexuée et cycle du développement [modifier]

La cellule est là aussi le vecteur de gènes et permet un brassage génétique au sein de la population
grâce aux processus cellulaires que sont la méiose et la fécondation.
La reproduction sexuée est caractéristique des eucaryotes, mais il existe des mécanismes de
brassage génétique chez les procaryotes.

La mort cellulaire : la cellule au service de l'organisme et des gènes [modifier]

La cellule ayant reçu un signal de son environnement va exprimer un programme entraînant sa mort
(l'apoptose étant un de ces mécanismes) : ce phénomène est nécessaire au développement des organismes
pluricellulaires ; autant chez les végétaux (avec par exemple la mort des cellules formant le tube criblé), que
chez les animaux (lors de la mise en place de la main chez l'homme : on a initialement une main palmée, la
mort des cellules permet l'individualisation des doigts). Ce phénomène a aussi été découvert chez certaines
bactéries (la mort cellulaire permet de limiter le nombre de bactéries lorsque les ressources sont
La cellule, tant pour les êtres pluricellulaires que pour les unicellulaires, constitue une structure
vouée avant tout à permettre la reproduction de l'organisme et donc la transmission d'une structure de base
contenant un programme génétique. Ainsi, certains auteurs ont été amenés à formuler la théorie du gène
égoïste, considérant les organismes (et donc les cellules) comme de simples structures destinées à assurer
la transmission et la prolifération des gènes (le gène proliférant alors pour lui-même est qualifié d'égoïste).

Interdépendance cellulaire : de la cellule à l'organisme [modifier]

La cellule, en constant échange avec l'extérieur dépend entièrement de celui-ci. Elle dépend aussi et
surtout des autres cellules, à plusieurs niveaux :
Les êtres unicellulaires : la cellule « bonne à tout faire » [modifier]
Ici, l'être vivant ne comporte qu'une cellule : celle-ci doit donc assurer toutes les fonctions vitales (se
nourrir, intégrer et réagir aux variations du milieu, proliférer...)
La cellule est donc en quelque sorte autonome mais elle dépend tout de même des autres cellules
(rares sont les cellules ne prélevant que dans le milieu des composés exclusivement inorganiques).
Il peut donc exister une interdépendance cellulaire, même pour les êtres unicellulaires.

L'organisme pluricellulaire : une communauté de cellules interdépendantes [modifier]

Ses cellules sont totalement dépendantes du bon fonctionnement des autres cellules: chacune
d'entre elles, bien qu'ayant le même matériel génétique (à de rares exceptions près: les gamètes, les
lymphocytes par exemple), exprime un programme génétique particulier qui la maintient dans une voie de
différenciation (plus ou moins poussée). Cette spécialisation implique le fractionnement d'opérations
effectuées dans une seule cellule pour les unicellulaires : les cellules d'un même organisme s'organisent en
différentes structures (tissus organes systèmes....) réalisant des fonctions particulières. Ce fractionnement
des fonctions nécessite une coordination entre cellules d'où l'émergence de systèmes de communication
entre cellules.
On a donc une interdépendance forte au sein même de l'organisme qui se superpose à la
dépendance aux autres êtres vivants.
La cellule eucaryote: formation de communautés de cellules intracellulaires [modifier]
La théorie endosymbiotique (théorie démontrée en ce qui concerne les mitochondries et les
chloroplastes) énonce que les cellules eucaryotes se sont formées à partir d'une cellule procaryote ayant
phagocyté puis domestiqué des bactéries : celles-ci seraient à l'origine des mitochondries. L'invagination de
cyanobactéries aurait donné naissance aux chloroplastes.
La cellule eucaryote dérive donc de l'association symbiotique de bactéries qui sont devenues
totalement interdépendantes au point de former une seule et même unité structurale et fonctionnelle.

Les principales structures cellulaires [modifier]

Il existe deux types fondamentaux de cellules selon qu'elles possèdent ou non un noyau :
• les procaryotes dont l'ADN est libre dans le cytoplasme (les bactéries, par exemple). Ils
comprennent les eubactéries et les archéobactéries ;
• les eucaryotes qui ont une organisation complexe, de nombreux organites et dont le noyau
est entouré d'une membrane nucléaire.
Principales différences entre les cellules procaryotes et eucaryotes
Procaryotes Eucaryotes

protistes, champignons, plantes,

représentants bactéries, archées

Taille typique ~ 1-10 µm ~ 10-100 µm

nucléoïde; pas de véritable
Type de noyau vrai noyau avec une enveloppe

circulaire (chromosome), molécules linéaires

ADN avec des protéines HU pour (chromosomes) avec des protéines
eubactéries histone
synthèse d'ARN dans le noyau
ARN/synthèse des
couplé au cytoplasme synthèse de protéines dans le

Ribosomes 23S+16S+5S 28S+18S+5,8S+5S

très structuré par des

très peu de structures membranes intracellulaires et un

Mouvement de la
flagelle fait de flagelline flagelle et cils fait de tubuline

Métabolisme anaérobie ou aérobie habituellement aérobie

Mitochondries aucune de une à plusieurs douzaines

Chloroplastes aucun dans les algues et les plantes


cellules isolées, colonies,

habituellement des cellules
Organisation organismes complexes avec des
cellules spécialisées

Mitose (multiplication conforme

Division de la
division simple de la cellule)
Méiose (formation de gamètes)

Eubactéries [modifier]
• Le cytoplasme des procaryotes (le contenu de la cellule) est diffus et granulaire, du fait des
ribosomes (complexe macromoléculaire responsable de la synthèse des protéines).
• La membrane plasmique isole l'intérieur de la cellule de son environnement, et sert de filtre
et de porte de communication.
• Il y a souvent [1] une paroi cellulaire. Elle est formée de peptidoglycane chez les
eubactéries, et joue le rôle de barrière supplémentaire contre les forces extérieures. Elle empêche
également la cellule d'éclater sous la pression osmotique dans un environnement hypotonique.
• L'ADN des procaryotes se compose d'une molécule circulaire super enroulée. Bien que sans
véritable noyau, l'ADN est toutefois condensé en un nucléoïde.
Les procaryotes peuvent posséder un ADN extra-chromosomal, organisé en molécules circulaires
appelées plasmides. Ils peuvent avoir des fonctions supplémentaires, telles que la résistance aux
antibiotiques. Certains procaryotes ont un flagelle leur permettant de se déplacer activement, plutôt que de
dériver passivement.

Spécificités des archées [modifier]

Les archéobactéries (archaea) sont considérées comme similaires à certains des premiers
organismes qui existèrent sur Terre. On les rencontre notamment dans des milieux extrêmes (elles sont
souvent appelées extrémophiles), tels que geysers, monts hydrothermaux, les fonds abyssaux. Certaines
peuvent résister à des pressions et des températures extrêmes, et avoir un métabolisme basé sur le
méthane ou le soufre.

Cellule eucaryote [modifier]

Organisation d'une cellule animale eucaryote typique.
1. Nucléole

2. Noyau

3. Ribosome

4. Vésicule

5. Réticulum
endoplasmique rugueux
6. Appareil de

7. Microtubule

8. Réticulum
endoplasmique lisse

9. Mitochondrie

10. Lysosome

11. Cytoplasme
(rempli par le cytosol)


13. Centrosome
• Le cytoplasme n'est pas aussi granulaire que celui des procaryotes, puisque la majeure
partie de ses ribosomes sont rattachés au réticulum endoplasmique.
• La membrane plasmique ressemble, dans sa fonction, à celle des procaryotes, avec
quelques différences mineures dans sa configuration.
• La paroi cellulosique, quand elle existe (végétaux), est composée de polysaccharides,
principalement la cellulose.
• L'ADN des eucaryotes est organisé en une ou plusieurs molécules linéaires. Ces molécules
se condensent en s'enroulant autour d'histones lors de la division cellulaire. Tous les chromosomes
de l'ADN sont stockés dans le noyau, séparés du cytoplasme par une membrane. Les eucaryotes ne
possèdent pas de plasmides : seuls quelques organites peuvent contenir de l'ADN.
• Certaines cellules eucaryotes peuvent devenir mobiles, en utilisant un cil ou un flagelle
(spermatozoïde par exemple). Leur flagelle est plus évolué que celui des procaryotes.
Les eucaryotes contiennent plusieurs organites. Ce sont des compartiments cellulaires baignant
dans le hyaloplasme. Ils sont délimités par une membrane plasmique (simple, double ou triple) et possèdent
des fonctions spécifiques.
• Le réticulum endoplasmique (RE) est une extension de la membrane du noyau. Il est divisé
en RE lisse (REL) et RE rugueux (RER) (parfois appelé RE granuleux REG), en fonction de son
apparence au microscope. La surface du RE rugueux est couverte de ribosomes qui insèrent les
protéines néosynthétisées dans le RE. Du RE, les protéines sont transportées vers l'appareil de
Golgi grâce à des vésicules.
• L'appareil de Golgi est le lieu de transformation finale des protéines. La glycosylation (ajout
de chaînes glucidiques complexes) se réalise à ce niveau.
• Les mitochondries jouent un rôle important dans le métabolisme de la cellule. Elles
contiennent leur propre génome (l'ADN mitochondrial). C'est là que se déroulent la respiration
cellulaire et la fabrication de l'énergie, l'ATP (Adénosine TriPhosphate). Cette énergie est
indispensable aux réactions métaboliques.
• Le cytosquelette permet à la cellule de conserver sa forme (Tenségrité) et de se mouvoir. Il
est également important lors de la division cellulaire, et dans le système de transport intracellulaire.
• Les plastes sont présents dans les plantes et les algues. Les plus connus sont les
chloroplastes, dans les cellules d'organismes photosynthétiques, qui convertissent l'énergie
lumineuse du Soleil en énergie chimique utilisée pour fabriquer des sucres à partir de dioxyde de
carbone (phase sombre de la photosynthèse). Ils possèdent également leur propre génome. Ils sont
le fruit de l'endosymbiose d'une cyanobactéries.
• Chez les plantes, les algues et les champignons, la cellule est encerclée par une paroi
cellulaire pectocellulosique qui fournit un squelette à l'organisme[2]. Des dépositions de composés
tels que la subérine ou la lignine modulent les propriétés physico-chimiques de la paroi, la rendant
plus solide ou plus imperméable, par exemple.
Certains eucaryotes unicellulaires peuvent former des structures multicellulaires. Ces colonies
consistent soit en des groupes de cellules identiques, capables de rester en vie une fois séparées de la
colonie principale (par exemple, les champignons), soit en des groupes de cellules spécialisées
Les méthodes d'étude de la cellule in vitro [modifier]

Microscopie [modifier]
La microscopie optique (résolution de +/- 0,25µm en lumière visible) permet l'observation de la
structure des cellules eucaryotes.
La microscopie électronique (résolution de quelques Angströms) révèle l'ultrastructure de celles-ci et
permet une observation plus poussée de la structure des cellules procaryotes comme eucaryotes.

Marquage de molécules [modifier]

Pour étudier l'organisation subcellulaire des cellules au microscope, les tissus peuvent, en fonction
de la méthode choisie, être vivants, ce qui permet une observation dynamique, ou fixés et préparés en
coupes histologiques, ce qui permet en général une observation plus précise, mais figée et ponctuelle.
Localisation subcellulaire par l'utilisation de gènes rapporteurs tels que la GFP (green fluorescing
protein) et la luciférase, par immunocytochimie, ou grâce à des molécules radioactives.
Différentes colorations, vitales ou non, permettent l'observation des structures au microscope
optique : rouge neutre pour les vacuoles, violet dahlia ou cristal pour le noyau...
Étude des constituants cellulaires [modifier]
Isolement de structures: par choc osmotique, ou grâce à des détergents puis par centrifugation.
Purification de protéines: par dialyse, chromatographie, électrophorèse....

Numération des cellules [modifier]

Il est fréquent de devoir compter le nombre de cellules vivantes dans une boîte de culture et de le
comparer au nombre de cellules total, par exemple pour déterminer la toxicité d'un produit. L'une de ces
méthodes de numération est réalisée grâce au test MTT.

Notes et références [modifier]

1. ↑ pas chez les mycoplasmes, par exemple
2. ↑ Geoffrey M. Cooper, La cellule : une approche moléculaire, De Boeck Université, 1999, p.

Voir aussi [modifier]

• Voir la vidéo "the inner life of the cell" faite par BioVision (harvard)
• Voir la vidéo sur la cellule dans l'encyclopédie médicale Vulgaris
Sur les autres projets Wikimédia :
• « Cellule (biologie) », sur Wikimedia Commons (ressources multimédia)
• « Cellule (biologie) », sur le Wiktionnaire (dictionnaire universel)
Liens internes [modifier]
• Biologie cellulaire
• Cellule souche
• Culture cellulaire
• Cellule végétale
• Respiration cellulaire
• Unicellulaire

Références externes [modifier]

• (fr) : le site de la Science du vivant
• (fr) Dossier Sagascience du CNRS : la cellule animale
• (fr) Qu'est ce qu'une cellule : La cellule en microcinéma
• (fr) Schéma détaillé d'une cellule animale.
• (fr) Schéma détaillé d'une cellule végétale.
• (fr) Exploration 3D d'une cellule :,
• (de) Cell Biology - Graphics
• biologie cellulaire des molécules aux organismes, Jean Claude Callen

• Portail de la biologie cellulaire et moléculaire


Introduction to evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is intended as an accessible, non-technical introduction to the subject. For the main
encyclopedia article, see Evolution.
Artist's depiction of a T. rex.
Natural selection does not lead to
perfection; dramatic changes in the
environment often lead to mass
extinctions, as in the case of the dinosaurs
nearly 65 million years ago.

Life forms reproduce to make


The offspring differs from the

parent in minor random ways.

If the differences are helpful, the

offspring is more likely to survive and

This means that more offspring in

the next generation will have the helpful

These differences accumulate

resulting in changes within the population.

Over time, populations branch off

to become new species as they become
geographically separated and genetically

This process is responsible for the

many diverse life forms in the world today.
Haeckel's Paleontological Tree of
Vertebrates (c. 1879).
The evolutionary history of species has
been described as a "tree", with many
branches arising from a single trunk. While
Haeckel's tree is somewhat outdated, it
illustrates clearly the principles that more
complex modern reconstructions can
Evolution is the process of change in all forms of life over generations, and evolutionary biology is the
study of how evolution occurs.
The biodiversity of life evolves by means of natural selection, mutations and genetic drift. The
principles of natural selection are based on three factual observations. First, every individual is supplied with
hereditary material in the form of genes that are received from their parents; then, passed on to their
offspring. Second, organisms tend to produce more offspring than the environment can support. Third, there
are variations among offspring as a consequence of either the random introduction of new genes via
mutations or reshuffling of existing genes during sexual reproduction.[1][2][3]
Natural selection will occur when these repeatedly observed facts of nature (heredity, overproduction
of offspring, and variation) hold true. Natural selection means individuals do not have equal chances of
reproductive success. As a consequence, some individuals produce more offspring and thus have a higher
degree of fitness. Traits that ensure organisms are better adapted to their living conditions become more
common in descendant populations.[2][3] For this reason, populations will never remain exactly the same
over successive generations. The forces of evolution are most evident when populations become isolated,
either through geographic distance or by mechanisms that prevent genetic exchange. Over time, isolated
populations can branch off into new species.[4][5] This is the basic premise behind evolutionary theory that
explains the origins of new species by means of natural selection.[2]
Random genetic drift describes another natural process that regulates the evolution of minor
mutations in the genes, leading to changes in allele frequencies over time. These smaller mutations occur
with regular frequency in and among populations. The vast majority of genetic mutations neither assist,
change the appearance of, nor bring harm to individuals. These mutated genes are neutrally sorted among
populations and survive across generations by chance alone. When species migrate they carry different
genetic varieties to different places. When organisms mate they exchange genetic material and new
individuals are born.
The outward expression of each unique genetic mixture in different environments, the phenotype,
creates a diversity of traits that can be measured and observed as individuals grow and develop. Physical
and behavioural traits are regulated by environmental conditions and, like clay, they are malleable as
individuals interact and respond to ever changing environmental and ecological situations. In contrast to
genetic drift, natural selection is not a random process because it acts on traits that become adapted for their
functional utilities that are necessary for survival.[6] Natural selection and random genetic drift are forever
constant and dynamic parts of life. More than 99.9% of all species have become extinct since life began over
3,500 million years ago. Evolution is more death than survival and over time this has shaped the branching
structure in the tree of life.[7]
The modern understanding of evolution began with the 1859 publication of Charles Darwin's On the
Origin of Species. In addition, Gregor Mendel's work with plants helped to explain the hereditary patterns of
genetics.[8] Fossil discoveries in paleontology, advances in population genetics and a global network of
scientific research have provided further details into the mechanisms of evolution. Scientists now have a
good understanding of the origin of new species (speciation) and have observed the speciation process in
the laboratory and in the wild. Evolution is the principal theory that biologists use to understand life and is
used in many disciplines, including medicine, psychology, conservation biology, anthropology, forensics,
agriculture and other social-cultural applications.
• 1 Darwin's idea: evolution by natural selection
• 2 Source of variation
• 3 Modern synthesis
• 4 Evidence for evolution
• 4.1 Fossil record
• 4.2 Comparative anatomy
• 4.2.1 Taxonomy
• 4.2.2 Embryology
• 4.2.3 Vestigial structures
• 4.2.4 Convergent evolution
• 4.3 Molecular biology
• 4.4 Co-evolution
• 4.5 Artificial selection
• 5 Species
• 6 Different views on the mechanism of evolution
• 6.1 Rate of change
• 6.2 Unit of change
• 7 Summary
• 8 See also
[edit] Darwin's idea: evolution by natural selection
Further information: Common descent
In the 19th century, natural history collections and museums were a popular pastime. The European
expansion and naval expeditions employed naturalists and curators of grand museums showcasing
preserved and live specimens of the varieties of life. Charles Darwin was an English graduate who was
educated and trained in the disciplines of natural history science. Such natural historians would collect,
catalogue, describe and study the vast collections of specimens stored and managed by curators at these
museums. Charles Darwin served as a ship's naturalist on board the HMS Beagle, assigned to a five-year
research expedition around the world. During his voyage, Darwin observed and collected an abundance of
organisms, being very interested in the diverse forms of life along the coasts of South America and the
neighboring Galapagos Islands.[9][10]
Charles Darwin gained extensive experience as he collected and studied the natural history of life
forms from distant places. Through his studies, Darwin formulated the idea that each species had developed
from ancestors with similar features. In 1838, he described how a process he called natural selection would
make this happen.[11]
Darwin's idea of how evolution works relied on the following observations:[12]
• If all the individuals of a species reproduced successfully, the population of that species
would increase uncontrollably.
• Populations tend to remain about the same size from year to year.
• Environmental resources are limited.
• No two individuals in a given species are exactly alike.
• Much of this variation in a population can be passed on to offspring.

Charles Darwin proposed the theory of evolution by natural selection.

Darwin noted that orchids exhibited a variety of complex adaptations to ensure pollination; all derived
from basic floral parts.
Darwin deduced that since organisms produce more offspring than their environment could possibly
support, there must be a competitive struggle for survival—only a few individuals can survive out of each
generation. Darwin realized that it was not chance alone that determined survival. Instead, survival depends
on the traits of each individual and if these traits aid or hinder survival and reproduction. Well-adapted, or
"fit", individuals are likely to leave more offspring than their less well-adapted competitors. Darwin realized
that the unequal ability of individuals to survive and reproduce could cause gradual changes in the
population. Traits that help an organism survive and reproduce would accumulate over generations. On the
other hand, traits that hinder survival and reproduction would disappear. Darwin used the term natural
selection to describe this process.[13]
Natural selection is commonly equated with survival of the fittest, but this expression originated in
Herbert Spencer's Principles of Biology in 1864, after Charles Darwin published his original works. Survival
of the fittest describes the process of natural selection incorrectly, because natural selection is not only about
survival and it is not always the fittest that survives.[14]
Observations of variations in animals and plants formed the basis of the theory of natural selection.
For example, Darwin observed that orchids and insects have a close relationship that allows the pollination of
the plants. He noted that orchids have a variety of structures that attract insects, so that pollen from the
flowers gets stuck to the insects’ bodies. In this way, insects transport the pollen from a male to a female
orchid. In spite of the elaborate appearance of orchids, these specialized parts are made from the same
basic structures that make up other flowers. In Fertilisation of Orchids Darwin proposed that the orchid
flowers did not represent the work of an ideal engineer, but were adapted from pre-existing parts, through
natural selection.[15]
Darwin was still researching and experimenting with his ideas on natural selection when he received
a letter from Alfred Wallace describing a theory very similar to his own. This led to an immediate joint
publication of both theories. Both Wallace and Darwin saw the history of life like a family tree, with each fork
in the tree’s limbs being a common ancestor. The tips of the limbs represented modern species and the
branches represented the common ancestors that are shared amongst many different species. To explain
these relationships, Darwin said that all living things were related, and this meant that all life must be
descended from a few forms, or even from a single common ancestor. He called this process descent with
Darwin published his theory of evolution by natural selection in On the Origin of Species in 1859. His
theory means that all life, including humanity, is a product of continuing natural processes. The implication
that all life on earth has a common ancestor has met with objections from some religious groups who believe
even today that the different types of life are due to special creation.[16] Their objections are in contrast to
the level of support for the theory by more than 99 percent of those within the scientific community today.[17]

[edit] Source of variation

Darwin’s theory of natural selection laid the groundwork for modern evolutionary theory, and his
experiments and observations showed that the organisms in populations varied from each other, that some
of these variations were inherited, and that these differences could be acted on by natural selection.
However, he could not explain the source of these variations. Like many of his predecessors, Darwin
mistakenly thought that heritable traits were a product of use and disuse, and that features acquired during
an organism's lifetime could be passed on to its offspring. He looked for examples, such as large ground
feeding birds getting stronger legs through exercise, and weaker wings from not flying until, like the ostrich,
they could not fly at all.[18] This misunderstanding was called the inheritance of acquired characters and was
part of the theory of transmutation of species put forward in 1809 by Jean-Baptiste Lamarck. In the late 19th
century this theory became known as Lamarckism. Darwin produced an unsuccessful theory he called
pangenesis to try to explain how acquired characteristics could be inherited. In the 1880s August
Weismann's experiments indicated that changes from use and disuse could not be inherited, and
Lamarckism gradually fell from favor.[19]
The missing information needed to help explain how new features could pass from a parent to its
offspring was provided by the pioneering genetics work of Gregor Mendel. Mendel’s experiments with several
generations of pea plants demonstrated that inheritance works by separating and reshuffling hereditary
information during the formation of sex cells and recombining that information during fertilization. This is like
mixing different hands of cards, with an organism getting a random mix of half of the cards from one parent,
and half of the cards from the other. Mendel called the information factors; however, they later became
known as genes. Genes are the basic units of heredity in living organisms. They contain the information that
directs the physical development and behavior of organisms.
Genes are made of DNA, a long molecule that carries information. This information is encoded in the
sequence of nucleotides in the DNA, just as the sequence of the letters in words carries information on a
page. The genes are like short instructions built up of the "letters" of the DNA alphabet. Put together, the
entire set of these genes gives enough information to serve as an "instruction manual" of how to build and
run an organism. The instructions spelled out by this DNA alphabet can be changed, however, by mutations,
and this may alter the instructions carried within the genes. Within the cell, the genes are carried in
chromosomes, which are packages for carrying the DNA, with the genes arranged along them like beads on
a string. It is the reshuffling of the chromosomes that results in unique combinations of genes in offspring.
Although such mutations in DNA are random, natural selection is not a process of chance: the
environment determines the probability of reproductive success. The end products of natural selection are
organisms that are adapted to their present environments. Natural selection does not involve progress
towards an ultimate goal. Evolution does not necessarily strive for more advanced, more intelligent, or more
sophisticated life forms.[20] For example, fleas (wingless parasites) are descended from a winged, ancestral
scorpionfly, and snakes are lizards that no longer require limbs - although pythons still grow tiny structures
that are the remains of their ancestor's hind legs.[21][22] Organisms are merely the outcome of variations
that succeed or fail, dependent upon the environmental conditions at the time.
Rapid environmental changes typically cause extinctions.[23] Of all species that have existed on
Earth, 99.9 percent are now extinct.[24] Since life began on Earth, five major mass extinctions have led to
large and sudden drops in the variety of species. The most recent, the Cretaceous–Tertiary extinction event,
occurred 65 million years ago, and has attracted more attention than all others because it killed the

[edit] Modern synthesis

Further information: Modern evolutionary synthesis
The modern evolutionary synthesis was the outcome of a merger of several different scientific fields
into a cohesive understanding of evolutionary theory. In the 1930s and 1940s, efforts were made to merge
Darwin's theory of natural selection, research in heredity, and understandings of the fossil records into a
unified explanatory model.[26] The application of the principles of genetics to naturally occurring populations,
by scientists such as Theodosius Dobzhansky and Ernst Mayr, advanced understanding of the processes of
evolution. Dobzhansky's 1937 work Genetics and the Origin of Species was an important step in bridging the
gap between genetics and field biology. Mayr, on the basis of an understanding of genes and direct
observations of evolutionary processes from field research, introduced the biological species concept, which
defined a species as a group of interbreeding or potentially interbreeding populations that are reproductively
isolated from all other populations. The paleontologist George Gaylord Simpson helped to incorporate fossil
research, which showed a pattern consistent with the branching and non-directional pathway of evolution of
organisms predicted by the modern synthesis.
The modern synthesis emphasizes the importance of populations as the unit 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 evolve as an accumulation of small changes over long periods of time.
[edit] Evidence for evolution

During the voyage of the Beagle, naturalist Charles Darwin collected fossils in South America, and
found fragments of armor which he thought were like giant versions of the scales on the modern armadillos
living nearby. On his return, the anatomist Richard Owen showed him that the fragments were from gigantic
extinct glyptodons, related to the armadillos. This was one of the patterns of distribution that helped Darwin to
develop his theory.[11]
Further information: Evidence of common descent
Scientific evidence for evolution comes from many aspects of biology, and includes fossils,
homologous structures, and molecular similarities between species' DNA.

[edit] Fossil record

Research in the field of paleontology, the study of fossils, supports the idea that all living organisms
are related. Fossils provide evidence that accumulated changes in organisms over long periods of time have
led to the diverse forms of life we see today. A fossil itself reveals the organism's structure and the
relationships between present and extinct species, allowing paleontologists to construct a family tree for all of
the life forms on earth.[27]
Modern paleontology began with the work of Georges Cuvier (1769–1832). Cuvier noted that, in
sedimentary rock, each layer contained a specific group of fossils. The deeper layers, which he proposed to
be older, contained simpler life forms. He noted that many forms of life from the past are no longer present
today. One of Cuvier’s successful contributions to the understanding of the fossil record was establishing
extinction as a fact. In an attempt to explain extinction, Cuvier proposed the idea of “revolutions” or
catastrophism in which he speculated that geological catastrophes had occurred throughout the earth’s
history, wiping out large numbers of species.[28] Cuvier's theory of revolutions was later replaced by
uniformitarian theories, notably those of James Hutton and Charles Lyell who proposed that the earth’s
geological changes were gradual and consistent.[29] However, current evidence in the fossil record supports
the concept of mass extinctions. As a result, the general idea of catastrophism has re-emerged as a valid
hypothesis for at least some of the rapid changes in life forms that appear in the fossil records.
A very large number of fossils have now been discovered and identified. These fossils serve as a
chronological record of evolution. The fossil record provides examples of transitional species that
demonstrate ancestral links between past and present life forms.[30] One such transitional fossil is
Archaeopteryx, an ancient organism that had the distinct characteristics of a reptile (such as a long, bony tail
and conical teeth) yet also had characteristics of birds (such as feathers and a wishbone). The implication
from such a find is that modern reptiles and birds arose from a common ancestor.[31]

[edit] Comparative anatomy

Further information: Convergent evolution and Divergent evolution
The comparison of similarities between organisms of their form or appearance of parts, called their
morphology, has long been a way to classify life into closely related groups. This can be done by comparing
the structure of adult organisms in different species or by comparing the patterns of how cells grow, divide
and even migrate during an organism's development.

[edit] Taxonomy
Taxonomy is the branch of biology that names and classifies all living things. Scientists use
morphological and genetic similarities to assist them in categorizing life forms based on ancestral
relationships. For example, orangutans, gorillas, chimpanzees, and humans all belong to the same
taxonomic grouping referred to as a family – in this case the family called Hominidae. These animals are
grouped together because of similarities in morphology that come from common ancestry (called homology).
Strong evidence for evolution comes from the analysis of homologous structures: structures in
different species that no longer perform the same task but which share a similar structure.[33] Such is the
case of the forelimbs of mammals. The forelimbs of a human, cat, whale, and bat all have strikingly similar
bone structures. However, each of these four species' forelimbs performs a different task. The same bones
that construct a bat's wings, which are used for flight, also construct a whale's flippers, which are used for
swimming. Such a "design" makes little sense if they are unrelated and uniquely constructed for their
particular tasks. The theory of evolution explains these homologous structures: all four animals shared a
common ancestor, and each has undergone change over many generations. These changes in structure
have produced forelimbs adapted for different tasks.[34]

[edit] Embryology
In some cases, anatomical comparison of structures in the embryos of two or more species provides
evidence for a shared ancestor that may not be obvious in the adult forms. As the embryo develops, these
homologies can be lost to view, and the structures can take on different functions. Part of the basis of
classifying the vertebrate group (which includes humans), is the presence of a tail (extending beyond the
anus) and pharyngeal slits. Both structures appear during some stage of embryonic development but are not
always obvious in the adult form.[35]
Because of the morphological similarities present in embryos of different species during
development, it was once assumed that organisms re-enact their evolutionary history as an embryo. It was
thought that human embryos passed through an amphibian then a reptilian stage before completing their
development as mammals. Such a re-enactment, (often called Recapitulation theory), is not supported by
scientific evidence. What does occur, however, is that the first stages of development are similar in broad
groups of organisms.[36] At very early stages, for instance, all vertebrates appear extremely similar, but do
not exactly resemble any ancestral species. As development continues, specific features emerge from this
basic pattern.

[edit] Vestigial structures

Homology includes a unique group of shared structures referred to as vestigial structures. Vestigial
refers to anatomical parts that are of minimal, if any, value to the organism that possesses them. These
apparently illogical structures are remnants of organs that played an important role in ancestral forms. Such
is the case in whales, which have small vestigial bones that appear to be remnants of the leg bones of their
ancestors which walked on land.[37] Humans also have vestigial structures, including the ear muscles, the
wisdom teeth, the appendix, the tail bone, body hair (including goose bumps), and the semilunar fold in the
corner of the eye.[38]

[edit] Convergent evolution

The bird and the bat wing are examples of convergent evolution.
A bat is a mammal and its forearm bones have been adapted for flight.
Anatomical comparisons can be misleading, as not all anatomical similarities indicate a close
relationship. Organisms that share similar environments will often develop similar physical features, a
process known as convergent evolution. Both sharks and dolphins have similar body forms, yet are only
distantly related – sharks are fish and dolphins are mammals. Such similarities are a result of both
populations being exposed to the same selective pressures. Within both groups, changes that aid swimming
have been favored. Thus, over time, they developed similar appearances (morphology), even though they
are not closely related.[39]
[edit] Molecular biology
A section of DNA
Every living organism (with the possible exception of RNA viruses) contains molecules of DNA,
which carries genetic information. Genes are the pieces of DNA that carry this information, and they influence
the properties of an organism. Genes determine an individual's general appearance and to some extent their
behavior. If two organisms are closely related, their DNA will be very similar.[40] On the other hand, the more
distantly related two organisms are, the more differences they will have. For example, brothers are closely
related and have very similar DNA, while cousins share a more distant relationship and have far more
differences in their DNA. Similarities in DNA are used to determine the relationships between species in
much the same manner as they are used to show relationships between individuals. For example, comparing
chimpanzees with gorillas and humans shows that there is as much as a 96 percent similarity between the
DNA of humans and chimps. Comparisons of DNA indicate that humans and chimpanzees are more closely
related to each other than either species is to gorillas.[41][42]
The field of molecular systematics focuses on measuring the similarities in these molecules and
using this information to work out how different types of organisms are related through evolution. These
comparisons have allowed biologists to build a relationship tree of the evolution of life on earth.[43] They
have even allowed scientists to unravel the relationships between organisms whose common ancestors lived
such a long time ago that no real similarities remain in the appearance of the organisms.

[edit] Co-evolution
Co-evolution is a process in which two or more species influence the evolution of each other. All
organisms are influenced by life around them; however, in co-evolution there is evidence that genetically
determined traits in each species directly resulted from the interaction between the two organisms.[40]
An extensively documented case of co-evolution is the relationship between Pseudomyrmex, a type
of ant, and the acacia, a plant that the ant uses for food and shelter. The relationship between the two is so
intimate that it has led to the evolution of special structures and behaviors in both organisms. The ant
defends the acacia against herbivores and clears the forest floor of the seeds from competing plants. In
response, the plant has evolved swollen thorns that the ants use as shelter and special flower parts that the
ants eat.[44] Such co-evolution does not imply that the ants and the tree choose to behave in an altruistic
manner. Rather, across a population small genetic changes in both ant and tree benefited each. The benefit
gave a slightly higher chance of the characteristic being passed on to the next generation. Over time,
successive mutations created the relationship we observe today.

[edit] Artificial selection

The results of artificial selection: a Chihuahua mix and a Great Dane.
Artificial selection is the controlled breeding of domestic plants and animals. Humans determine
which animal or plant will reproduce and which of the offspring will survive; thus, they determine which genes
will be passed on to future generations. The process of artificial selection has had a significant impact on the
evolution of domestic animals. For example, people have produced different types of dogs by controlled
breeding. The differences in size between the Chihuahua and the Great Dane are the result of artificial
selection. Despite their dramatically different physical appearance, they and all other dogs evolved from a
few wolves domesticated by humans in what is now China less than 15,000 years ago.[45]
Artificial selection has produced a wide variety of plants. In the case of maize (corn), recent genetic
evidence suggests that domestication occurred 10,000 years ago in central Mexico.[46] Prior to
domestication, the edible portion of the wild form was small and difficult to collect. Today The Maize Genetics
Cooperation • Stock Center maintains a collection of more than 10,000 genetic variations of maize that have
arisen by random mutations and chromosomal variations from the original wild type.[47]
In artificial selection the new breed or variety that emerges is the one with random mutations
attractive to humans, while in natural selection the surviving species is the one with random mutations useful
to it in its non-human environment. In both natural and artificial selection the variations are a result of random
mutations, and the underlying genetic processes are essentially the same.[48] Darwin carefully observed the
outcomes of artificial selection in animals and plants to form many of his arguments in support of natural
selection.[49] Much of his book On the Origin of Species was based on these observations of the many
varieties of domestic pigeons arising from artificial selection. Darwin proposed that if humans could achieve
dramatic changes in domestic animals in short periods, then natural selection, given millions of years, could
produce the differences seen in living things today.
[edit] Species

There are numerous species of cichlids that demonstrate dramatic variations in morphology.
Further information: Species, Speciation, and Phylogenetics
Given the right circumstances, and enough time, evolution leads to the emergence of new species.
Scientists have struggled to find a precise and all-inclusive definition of species. Ernst Mayr (1904–2005)
defined a species as a population or group of populations whose members have the potential to interbreed
naturally with one another to produce viable, fertile offspring. (The members of a species cannot produce
viable, fertile offspring with members of other species).[50] Mayr's definition has gained wide acceptance
among biologists, but does not apply to organisms such as bacteria, which reproduce asexually.
be substantial, so that genetic exchange between the two populations is completely disrupted. In their
separate environments, the genetically isolated groups follow their own unique evolutionary pathways. Each
group will accumulate different mutations as well as be subjected to different selective pressures. The
accumulated genetic changes may result in separated populations that can no longer interbreed if they are
reunited.[13] Barriers that prevent interbreeding are either prezygotic (prevent mating or fertilization) or
postzygotic (barriers that occur after fertilization). If interbreeding is no longer possible, then they will be
considered different species.[51]
Usually the process of speciation is slow, occurring over very long time spans; thus direct
observations within human life-spans are rare. However speciation has been observed in present day
organisms, and past speciation events are recorded in fossils.[52][53][54] Scientists have documented the
formation of five new species of cichlid fishes from a single common ancestor that was isolated fewer than
5000 years ago from the parent stock in Lake Nagubago.[55] The evidence for speciation in this case was
morphology (physical appearance) and lack of natural interbreeding. These fish have complex mating rituals
and a variety of colorations; the slight modifications introduced in the new species have changed the mate
selection process and the five forms that arose could not be convinced to interbreed.[56]
[edit] Different views on the mechanism of evolution

James Hutton
Stephen Jay Gould
Richard Dawkins
The theory of evolution is widely accepted among the scientific community, serving to link the diverse
specialty areas of biology.[17] Evolution provides the field of biology with a solid scientific base. The
significance of evolutionary theory is best described by the title of a paper by Theodosius Dobzhansky (1900–
1975), published in American Biology Teacher; "Nothing in Biology Makes Sense Except in the Light of
Evolution".[57] Nevertheless, the theory of evolution is not static. There is much discussion within the
scientific community concerning the mechanisms behind the evolutionary process. For example, the rate at
which evolution occurs is still under discussion. In addition, there are conflicting opinions as to which is the
primary unit of evolutionary change – the organism or the gene.
[edit] Rate of change
Two views exist concerning the rate of evolutionary change. Darwin and his contemporaries viewed
evolution as a slow and gradual process. Evolutionary trees are based on the idea that profound differences
in species are the result of many small changes that accumulate over long periods.
The view that evolution is gradual had its basis in the works of the geologist James Hutton (1726–
1797) and his theory called "gradualism". Hutton's theory suggests that profound geological change was the
cumulative product of a relatively slow continuing operation of processes which can still be seen in operation
today, as opposed to catastrophism which promoted the idea that sudden changes had causes which can no
longer be seen at work. A uniformitarian perspective was adopted for biological changes. Such a view can
seem to contradict the fossil record, which shows evidence of new species appearing suddenly, then
persisting in that form for long periods. The paleontologist Stephen Jay Gould (1941–2002) developed a
model that suggests that evolution, although a slow process in human terms, undergoes periods of relatively
rapid change over only a few thousand or million years, alternating with long periods of relative stability, a
model called "punctuated equilibrium" which explains the fossil record without contradicting Darwin's ideas.

[edit] Unit of change

A common unit of selection in evolution is the organism. Natural selection occurs when the
reproductive success of an individual is improved or reduced by an inherited characteristic, and reproductive
success is measured by the number of an individual's surviving offspring. The organism view has been
challenged by a variety of biologists as well as philosophers. Richard Dawkins (born 1941) proposes that
much insight can be gained if we look at evolution from the gene's point of view; that is, that natural selection
operates as an evolutionary mechanism on genes as well as organisms.[59] In his 1976 book The Selfish
Gene, he explains:
“ Individuals are not stable things, they are fleeting. Chromosomes too are
shuffled to oblivion, like hands of cards soon after they are dealt. But the cards
themselves survive the shuffling. The cards are the genes. The genes are not destroyed
by crossing-over; they merely change partners and march on. Of course they march on.
That is their business. They are the replicators and we are their survival machines.
When we have served our purpose we are cast aside. But genes are denizens of
geological time: genes are forever.[60] ”
Others view selection working on many levels, not just at a single level of organism or gene; for
example, Stephen Jay Gould called for a hierarchical perspective on selection.[61]

[edit] Summary
Several basic observations establish the theory of evolution, which explains the variety and
relationship of all living things. There are genetic variations within a population of individuals. Some
individuals, by chance, have features that allow them to survive and thrive better than their kind. The
individuals that survive will be more likely to have offspring of their own. The offspring might inherit the useful
neighbors in a particular environment. Fossils, the genetic code, and the peculiar distribution of life on earth
provide a record of evolution and demonstrate the common ancestry of all organisms, both living and long
dead. Evolution can be directly observed in artificial selection, the selective breeding for certain traits of
domestic animals and plants. The diverse breeds of cats, dogs, horses, and agricultural plants serve as
examples of evolution.
Although some groups raise objections to the theory of evolution, the evidence of observation and
experiments over a hundred years by thousands of scientists supports evolution.[16] The result of four billion
years of evolution is the diversity of life around us, with an estimated 1.75 million different species in
existence today.[5][62]

[edit] See also

Evolutionary biology portal


Books are collections of articles that can be downloaded or ordered in print.

• Creation-evolution controversy
• Evidence of common descent
• Evolution as theory and fact
• Level of support for evolution
• Misconceptions about evolution

[edit] Notes
1. ^ Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray. p. 1. . Related
earlier ideas were acknowledged in Darwin, Charles (1861). On the Origin of Species (3rd ed.).
London: John Murray. xiii.
2. ^ a b c Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard University
Press. pp. 1433. ISBN 0674006135, 9780674006133.
3. ^ a b Gregory, T. R. (2009). "Understanding Natural Selection: Essential Concepts and
Common Misconceptions". Evolution: Education and Outreach 2 (2): 156–175. doi:10.1007/s12052-
4. ^ "An introduction to evolution" (web resource). Understanding Evolution: your one-stop
source for information on evolution. The University of California Museum of Paleontology, Berkeley.
2008. Retrieved 2008-01-23
5. ^ a b Cavalier-Smith T (2006). "Cell evolution and Earth history: stasis and revolution". Philos
Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. doi:10.1098/rstb.2006.1842. PMID 16754610.
PMC 1578732. Retrieved 2008-01-24.
6. ^ Garvin-Doxas, K.; Klymkowsky, M. W. (2008). "Understanding Randomness and its Impact
on Student Learning: Lessons Learned from Building the Biology Concept Inventory (BCI)". CBE Life
Sci Educ. 7 (2): 227–233. doi:10.1187/cbe.07-08-0063. PMID 18519614. PMC 2424310.
7. ^ Raup, D. M. (1992). Extinction: bad genes or bad luck.. New York, W. W.: Norton and Co..
pp. 210. ISBN 978-0393309270.
8. ^ Rhee, Sue Yon (1999). "Gregor Mendel". Access Excellence. National Health Museum. Retrieved 2008-01-05.
9. ^ Farber, P. L. (2000). Finding Order in Nature: The Natualist Tradition from Linnaeus to E.
O. Wilson. Baltimore and London: The Johns Hopkins University Press. pp. 136. ISBN 0-8018-6389-
10.^ Watson, J. D. (2005). Darwin, The Indelible Stamp: The Evolution of an Idea . Philadelphia
and London: Running Press. pp. 1257. ISBN 13-978-0-7624-2136-7.
11.^ a b Eldredge, Niles (Spring 2006). "Confessions of a Darwinist". The Virginia Quarterly
Review: 32–53.
Retrieved 2008-01-23.
12.^ a b Wyhe, John van (2002). "Charles Darwin: gentleman naturalist". The Complete Work of
Charles Darwin Online. University of Cambridge. Retrieved
13.^ a b c Quammen, David (2004). "Was Darwin Wrong?". National Geographic Magazine.
National Geographic.
Retrieved 2007-12-23.
14.^ Futuyma, D. J. (2005). The Nature of Natural Selection. Ch. 8, pages 93-98 in Cracraft, J.
and Bybee R. W. (Eds.) Evolutionary Science and Society: Educating a New Generation. American
Institute of Biological Sciences.
15.^ Wyhe, John van (2002). "Fertilisation of Orchids". The Complete Works of Charles Darwin.
University of Cambridge. http://darwin- Retrieved 2008-01-07.
16.^ a b DeVries A (2004). "The enigma of Darwin". Clio Med 19 (2): 136–55. PMID 6085987.
17.^ a b Delgado, Cynthia (2006). "Finding the Evolution in Medicine". NIH Record (National
Institutes of Health).
Retrieved 2007-12-21.
18.^ (Darwin 1872, p. 108.) Effects of the increased Use and Disuse of Parts, as controlled by
Natural Selection
19.^ Ghiselin, Michael T. (September/October 1994). "Nonsense in schoolbooks: 'The Imaginary
Lamarck'". The Textbook Letter. The Textbook League.
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20.^ (Gould (a) 1981, p. 24)
21.^ Bejder L, Hall BK (2002). "Limbs in whales and limblessness in other vertebrates:
mechanisms of evolutionary and developmental transformation and loss". Evol. Dev. 4 (6): 445–58.
doi:10.1046/j.1525-142X.2002.02033.x. PMID 12492145.
22.^ Boughner JC, Buchtová M, Fu K, Diewert V, Hallgrímsson B, Richman JM (2007).
"Embryonic development of Python sebae - I: Staging criteria and macroscopic skeletal
morphogenesis of the head and limbs". Zoology (Jena) 110 (3): 212–30.
doi:10.1016/j.zool.2007.01.005. PMID 17499493.
23.^ Drummond, A; Strimmer, K (Jul 2001). Frequently Asked Questions About Evolution.
"Evolution Library". Bioinformatics (Oxford, England) (WGBH Educational Foundation) 17 (7): 662–3.
ISSN 1367-4803. PMID 11448888.
Retrieved 2008-01-23
24.^ "Roundtable: Mass Extinction". Evolution: a jouney into where we're from and where we're
going. WGBH Educational Foundation. 2001. Retrieved 2008-01-23 .
25.^ Bambach, R.K.; Knoll, A.H.; Wang, S.C. (December 2004). "Origination, extinction, and
mass depletions of marine diversity". Paleobiology 30 (4): 522–42. doi:10.1666/0094-
8373(2004)030<0522:OEAMDO>2.0.CO;2. Retrieved 2008-01-24
26.^ Committee on Defining and Advancing the Conceptual Basis of Biological Sciences (1989).
"The tangled web of biological science". The role of theory in advancing 21st Century
Biology:Catalyzing Transformation Research. National Research Council. Retrieved 2008-01-06.
27.^ "The Fossil Record - Life's Epic". The Virtual Fossil Museum. Retrieved 2007-08-31.
28.^ (Tattersall 1995, pp. 5–6)
29.^ (Lyell 1830, p. 76)
30.^ Committee on Revising Science and Creationism: A View from the National Academy of
Sciences, National Academy of Sciences and Institute of Medicine of the National Academies (2008).
"Science, Evolution, and Creationism". National Academy of Sciences. Retrieved 2008-01-06.
31.^ (Gould (b) 1995, p. 360)
32.^ (Diamond 1992, p. 16)
33.^ a b Drummond, A; Strimmer, K (Jul 2001). Glossary. "Evolution Library" (web resource).
Bioinformatics (Oxford, England) (WGBH Educational Foundation) 17 (7): 662–3. ISSN 1367-4803.
PMID 11448888. Retrieved 2008-01-
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34.^ (Mayr 2001, pp. 25–27)
35.^ (Weichert & Presch 1975, p. 8)
36.^ Miller, Kenneth (1997). "Haeckel and his Embryos". Evolution Resources. Retrieved 2007-09-02.
37.^ Theobald, Douglas (2004). "29+ Evidences for Macroevolution Part 2: Past History" (pdf).
TalkOrigins Archive. The TalkOrigins Foundation.
Retrieved 2008-01-27.
38.^ Johnson, George (2002). "Vestigial Structures" (web resource). The Evidence for Evolution.
'On Science' column in St. Louis Post Dispatch. Retrieved 2008-01-23 .
39.^ Johnson, George (2002). "Convergent and Divergent Evolution" (web resource). The
Evidence for Evolution. 'On Science' column in St. Louis Post Dispatch. Retrieved 2008-01-23 .
40.^ a b Kennedy, Donald; (Working group on teaching evolution) (1998). "Teaching about
evolution and the nature of science". Evolution and the nature of science. The National Academy of
Science. Retrieved 2007-12-30.
41.^ Lovgren, Stefan (2005-08-31). "Chimps, Humans 96 Percent the Same, Gene Study
Finds". National Geographic News. National Geographic. Retrieved
42.^ (Carroll, Grenier & Weatherbee 2000)
43.^ Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P (2006). "Toward
automatic reconstruction of a highly resolved tree of life". Science 311 (5765): 1283–87.
doi:10.1126/science.1123061. PMID 16513982.
44.^ Janzen, Daniel (1974). "Swollen-Thorn Acacias of Central America" (pdf). Smithsonian
Contributions to Biology. Smithsonian Institution. Retrieved 2007-08-31.
45.^ McGourty, Christine (2002-11-22). "Origin of dogs traced". BBC News. Retrieved 2007-12-14.
46.^ Hall, Hardy. "Transgene Escape: Are Traditioanl Corn Varieties In Mexico Threatened by
Transgenic Corn Crops". Scientific Creative Quarterly.
traditional-corn-varieties-in-mexico-threatened-by-transgenic-corn-crops/?. Retrieved 2007-12-14.
47.^ "The Maize Genetics Cooperation • Stock Center". National Plant Germplasm. U.S.
Department of Agriculture. 2006-06-21. Retrieved
48.^ Silverman, E. David (2002). "Better Books by Trial and Error.". Retrieved 2008-04-04.
49.^ Wilner A. (2006). "Darwin's artificial selection as an experiment". Stud Hist Philos Biol
Biomed Sci. 37 (1): 26–40. doi:10.1016/j.shpsc.2005.12.002. PMID 16473266.
50.^ (Mayr 2001, pp. 165–69)
51.^ Sulloway, Frank J (December 2005). "The Evolution of Charles Darwin". Smithsonian
Magazine. Smithsonian Institution. Retrieved 2007-
52.^ Jiggins CD, Bridle JR (2004). "Speciation in the apple maggot fly: a blend of vintages?".
Trends Ecol. Evol. (Amst.) 19 (3): 111–14. doi:10.1016/j.tree.2003.12.008. PMID 16701238.
53.^ Boxhorn, John (1995). "Observed Instances of Speciation". TalkOrigins Archive. Retrieved 2007-05-10.
54.^ Weinberg JR, Starczak VR, Jorg, D (1992). "Evidence for Rapid Speciation Following a
Founder Event in the Laboratory". Evolution (Evolution, Vol. 46, No. 4) 46 (4): 1214–20.
doi:10.2307/2409766. Retrieved 2008-01-24.
55.^ (Mayr 1970, p. 348)
56.^ (Mayr 1970)
57.^ "NCSE Resource". Cans and Can`ts of Teaching Evolution. National Center for Science
Education. 2001-02-13.
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58.^ Gould, Stephen Jay (1991). "Opus 200". Stephen Jay Gould Archive. Natural History. Retrieved 2007-08-31.
59.^ Wright, Sewall (September 1980). "Genic and Organismic Selection". Evolution (Evolution,
Vol. 34, No. 5) 34 (5): 825. doi:10.2307/2407990.
Z&size=LARGE&origin=JSTOR-enlargePage. Retrieved 2007-12-23.
60.^ (Dawkins 1976, p. 35)
61.^ Gould SJ, Lloyd EA (1999). "Individuality and adaptation across levels of selection: how
shall we name and generalize the unit of Darwinism?". Proc. Natl. Acad. Sci. U.S.A. 96 (21): 11904–
9. doi:10.1073/pnas.96.21.11904. PMID 10518549. PMC 18385. Retrieved 2008-01-18.
62.^ Sedjo, Roger (2007). "How many species are there?". Environmental Literacy Council. Retrieved 2008-01-05.

[edit] References
• Carroll, SB; Grenier, J; Weatherbee, SD (2000). From DNA to Diversity: Molecular Genetics
and the Evolution of Animal Design (2nd ed.). Oxford: Blackwell Publishing. ISBN 1-4051-1950-0
• Darwin, Charles (1872). The Origin of Species (6th ed.). London: John Murray. http://darwin-
• Dawkins, Richard (1976). The Selfish Gene (1st ed.). Oxford University Press. pp. 33.
ISBN 0192860925.
• Diamond, Jared (1992). The Third Chimpanzee: the evolution and future of the human
animal. New York: HarperCollins. ISBN 0060183071
• Gould (a), Stephen Jay (1981). The Panda's Thumb: More Reflections in Natural History .
New York: W.W, Norton & Company. ISBN 0393308197
• Gould (b), Stephen Jay (1995). Dinosaur in a Haystack. New York: Harmony Books.
ISBN 0517703939
• Lyell, Charles (1830). Principles of geology. New York: Penguin Books. ISBN 014043528X.
• Mayr, Ernst (1970). Populations, Species, and Evolution. Cambridge, MA: Belknap Press of
Harvard University Press. ISBN 0674690109
• Mayr, Ernst (2001). What evolution is. New York: Basic Books. ISBN 0-465-04425-5
• Tattersall, Ian (1995). The Fossil Trail: How We Know What We Think We Know About
Human Evolution. New York: Oxford University Press. ISBN 0195061012
• Weichert, Charles; Presch, William (1975). Elements of Chordate Anatomy. New York:
McGraw-Hill. ISBN 0070690081

[edit] Further reading

• Liam Neeson (narrator). (2001-11-20) (web resource). Evolution: a journey into where we're
from and where we're going. [DVD]. South Burlington, VT: WGBH Boston / PBS television series
Nova. ASIN B00005RG6J. Retrieved 2008-01-24. -
Age level: Grade 7+
• Horvitz, Leslie Alan (2002). The complete idiot's guide to evolution. Indianapolis: Alpha
Books. ISBN 0028642260.
• Charlesworth, Deborah; Charlesworth, Brian (2003). Evolution: a very short introduction.
Oxford: Oxford University Press. ISBN 0192802518.
• Sis, Peter (2003). The tree of life: a book depicting the life of Charles Darwin, naturalist,
geologist & thinker. New York: Farrar Straus Giroux. ISBN 0-374-45628-3.
• Thomson, Keith Stewart (2005). Fossils: a very short introduction. Oxford: Oxford University
Press. ISBN 0192805045.
• Greg Krukonis (2008). Evolution For Dummies (For Dummies (Math & Science)) . For
Dummies. ISBN 0-470-11773-7.
• Darwin, Charles (2008). Quammen, David. ed. On the Origin of Species: The Illustrated
Edition. Sterling. ISBN 1402756399
• Pallan, Mark (2009). The Rough Guide to Evolution. Rough Guides. ISBN 1858289467
• Zimmer, Carl (2009). The Tangled Bank: An Introduction to Evolution . Roberts and Company
Publishers. ISBN 0981519474
• Ellis, R. John (2010). How Science Works: Evolution. Springer. ISBN 9048131820

[edit] External links

• Brain, Marshall. "How Stuff Works: Evolution Library" (web resource). Retrieved 2008-01-24
• Carl Sagan. (2006-07-06) (Google video). Carl Sagan on evolution. [streaming video].
Google. Retrieved
• Carl Sagan. (2006-10-21) (Youtube video). Theory of Evolution Explained. [streaming video].
Youtube. Retrieved 2008-01-24.
• "Evolution Education Wiki: EvoWiki" (web resource). Retrieved 2008-
• "The Big Picture on Evolution (PDF)". The Big Picture Series. Wellcome Trust. January 2007.
ocument/wtd026042.pdf. Retrieved 2008-01-23
• "The Talk Origins Archive: Exploring the Creation/Evolution Controversy" (web resource). Retrieved 2008-01-24
• (web resource) Understanding Evolution: your one-stop source for information on evolution .
The University of California Museum of Paleontology, Berkeley. Retrieved 2008-01-24
• "University of Utah Genetics Learning Center animated tour of the basics of genetics" (web
resource). Retrieved 2008-01-
• "Introduction To Evolution" (web resource). Retrieved 2010-06-01

[edit] Related information

Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary Canalisation · Modularity · Phenotypic plasticity

biology (Evo-devo)

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

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Categories: Introductions | Biological evolution | Biology theories

• This page was last modified on 21 August 2010 at 20:55.


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Biology portal · v • d • e
Evolution is the change in the inherited traits of a population of organisms through successive
generations.[1] After a population splits into smaller groups, these groups evolve independently and may
eventually diversify into new species. Ultimately, life is descended from a common ancestry through a long
series of these speciation events, stretching back in a tree of life that has grown over the 3.5 billion years of
life on Earth. This is visible in anatomical, genetic and other likenesses between groups of organisms,
geographical distribution of related species, the fossil record and the recorded genetic changes in living
organisms over many generations. To distinguish from other uses of the word evolution, it is sometimes
termed biological evolution, genetic evolution or organic evolution.[2][3][4]
Evolution is the product of two opposing forces: processes that constantly introduce variation in traits,
and processes that make particular variants become more common or rare. A trait is a particular
characteristic, such as eye color, height, or a behavior, that is expressed when an organism's genes interact
with its environment. Genes vary within populations, so organisms show heritable differences (variation) in
their traits. The main cause of variation is mutation, which changes the sequence of a gene. Altered genes,
or alleles, are then inherited by offspring. There can sometimes also be transfer of genes between species.
Two main processes cause variants to become more common or rare in a population. One is natural
selection, through which traits that aid survival and reproduction become more common, while traits that
hinder survival and reproduction become more rare.[1][5] Natural selection occurs because only a few
individuals in each generation will survive, since resources are limited and organisms produce many more
offspring than their environment can support. Over many generations, mutations produce successive, small,
random changes in traits, which are then filtered by natural selection and the beneficial changes retained.
This adjusts traits so they become suited to an organism's environment: these adjustments are called
adaptations.[6] Not every trait, however, is an adaptation. Another cause of evolution is genetic drift, which
produces entirely random changes in how common traits are in a population. Genetic drift comes from the
role that chance plays in whether a trait will be passed on to the next generation.
Evolutionary biologists document the fact that evolution occurs, and also develop and test theories
that explain its causes. The study of evolutionary biology began in the mid-nineteenth century, when
research into the fossil record and the diversity of living organisms convinced most scientists that species
changed over time.[7][8] The mechanism driving these changes remained unclear until the theories of natural
selection were independently proposed by Charles Darwin and Alfred Wallace. In 1859, Darwin's seminal
work On the Origin of Species brought the new theories of evolution by natural selection to a wide audience,
[9] leading to the overwhelming acceptance of evolution among scientists.[10][11][12][13] In the 1930s,
Darwinian natural selection became understood in combination with Mendelian inheritance, forming the
modern evolutionary synthesis,[14] which connected the units of evolution (genes) and the mechanism of
evolution (natural selection). This powerful explanatory and predictive theory has become the central
organizing principle of modern biology, directing research and providing a unifying explanation for the history
and diversity of life on Earth.[11][12][15] Evolution is therefore applied and studied in fields as diverse as
agriculture, anthropology, conservation biology, ecology, medicine, paleontology, philosophy, and
psychology along with other specific topics in the previous listed fields.
• 1 History of evolutionary thought
• 2 Heredity
• 3 Variation
• 3.1 Mutation
• 3.2 Sex and recombination
• 3.3 Population genetics
• 3.4 Gene flow
• 4 Mechanisms
• 4.1 Natural selection
• 4.2 Genetic drift
• 5 Outcomes
• 5.1 Adaptation
• 5.2 Co-evolution
• 5.3 Co-operation
• 5.4 Speciation
• 5.5 Extinction
• 6 Evolutionary history of life
• 6.1 Origin of life
• 6.2 Common descent
[edit] History of evolutionary thought
For more details on this topic, see History of evolutionary thought.

Around 1854 Charles Darwin began writing out what became On the Origin of Species.
The scientific inquiry into the origin of species can be dated to at least the 6th century BCE, with the
Greek philosopher Anaximander.[16] Others who considered evolutionary ideas included the Greek
philosopher Empedocles, the Roman philosopher-poet Lucretius, the Afro-Arab biologist Al-Jahiz,[17] the
Persian philosopher Ibn Miskawayh, the Brethren of Purity,[18] and the Chinese philosopher Zhuangzi.[19]
As biological knowledge grew in the 18th century, evolutionary ideas were set out by a few natural
philosophers including Pierre Maupertuis in 1745 and Erasmus Darwin in 1796.[20] The ideas of the biologist
Jean-Baptiste Lamarck about transmutation of species influenced radicals, but were rejected by mainstream
scientists. Charles Darwin formulated his idea of natural selection in 1838 and was still developing his theory
in 1858 when Alfred Russel Wallace sent him a similar theory, and both were presented to the Linnean
Society of London in separate papers.[21] At the end of 1859, Darwin's publication of On the Origin of
Species explained natural selection in detail and presented evidence leading to increasingly wide acceptance
of the occurrence of evolution.
Debate about the mechanisms of evolution continued, and Darwin could not explain the source of the
heritable variations which would be acted on by natural selection. Like Lamarck, he thought that parents
passed on adaptations acquired during their lifetimes,[22] a theory which was subsequently dubbed
Lamarckism.[23] In the 1880s, August Weismann's experiments indicated that changes from use and disuse
were not heritable, and Lamarckism gradually fell from favour.[24][25] More significantly, Darwin could not
account for how traits were passed down from generation to generation. In 1865 Gregor Mendel found that
traits were inherited in a predictable manner.[26] When Mendel's work was rediscovered in 1900s,
disagreements over the rate of evolution predicted by early geneticists and biometricians led to a rift between
the Mendelian and Darwinian models of evolution.
Yet it was the rediscovery of Gregor Mendel's pioneering work on the fundamentals of genetics (of
which Darwin and Wallace were unaware) by Hugo de Vries and others in the early 1900s that provided the
impetus for a better understanding of how variation occurs in plant and animal traits. That variation is the
main fuel used by natural selection to shape the wide variety of adaptive traits observed in organic life. Even
though Hugo de Vries and other early geneticists rejected gradual natural selection, their rediscovery of and
subsequent work on genetics eventually provided a solid basis on which the theory of evolution stood even
more convincingly than when it was originally proposed.[27]
The apparent contradiction between Darwin's theory of evolution by natural selection and Mendel's
work was reconciled in the 1920s and 1930s by evolutionary biologists such as J.B.S. Haldane, Sewall
Wright, and particularly Ronald Fisher, who set the foundations for the establishment of the field of population
genetics. The end result was a combination of evolution by natural selection and Mendelian inheritance, the
modern evolutionary synthesis.[28] In the 1940s, the identification of DNA as the genetic material by Oswald
Avery and colleagues and the subsequent publication of the structure of DNA by James Watson and Francis
Crick in 1953, demonstrated the physical basis for inheritance. Since then, genetics and molecular biology
have become core parts of evolutionary biology and have revolutionized the field of phylogenetics.[14]
In its early history, evolutionary biology primarily drew in scientists from traditional taxonomically
oriented disciplines, whose specialist training in particular organisms addressed general questions in
evolution. As evolutionary biology expanded as an academic discipline, particularly after the development of
the modern evolutionary synthesis, it began to draw more widely from the biological sciences.[14] Currently
the study of evolutionary biology involves scientists from fields as diverse as biochemistry, ecology, genetics
and physiology, and evolutionary concepts are used in even more distant disciplines such as psychology,
medicine, philosophy and computer science. In the 21st century, current research in evolutionary biology
deals with several areas where the modern evolutionary synthesis may need modification or extension, such
as assessing the relative importance of various ideas on the unit of selection and evolvability and how to fully
incorporate the findings of evolutionary developmental biology.[29][30]

[edit] Heredity
Further information: Introduction to genetics, Genetics, and Heredity
DNA structure. Bases are in the center, surrounded by phosphate–sugar chains in a double helix.
Evolution in organisms occurs through changes in heritable traits – particular characteristics of an
organism. In humans, for example, eye color is an inherited characteristic and an individual might inherit the
"brown-eye trait" from one of their parents.[31] Inherited traits are controlled by genes and the complete set
of genes within an organism's genome is called its genotype.[32]
The complete set of observable traits that make up the structure and behavior of an organism is
called its phenotype. These traits come from the interaction of its genotype with the environment.[33] As a
result, many aspects of an organism's phenotype are not inherited. For example, suntanned skin comes from
the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's
children. However, some people tan more easily than others, due to differences in their genotype; a striking
example are people with the inherited trait of albinism, who do not tan at all and are very sensitive to
Heritable traits are passed from one generation to the next via DNA, a molecule that encodes genetic
information.[32] DNA is a long polymer composed of four types of bases. The sequence of bases along a
particular DNA molecule specify the genetic information, in a manner similar to a sequence of letters spelling
out a sentence. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the
DNA sequence.
Portions of a DNA molecule that specify a single functional unit are called genes; different genes
have different sequences of bases. Within cells, the long strands of DNA form condensed structures called
chromosomes. The specific location of a DNA sequence within a chromosome is known as a locus. If the
DNA sequence at a locus varies between individuals, the different forms of this sequence are called alleles.
DNA sequences can change through mutations, producing new alleles. If a mutation occurs within a gene,
the new allele may affect the trait that the gene controls, altering the phenotype of the organism.
However, while this simple correspondence between an allele and a trait works in some cases, most
traits are more complex and are controlled by multiple interacting genes.[35][36] The study of such complex
traits is a major area of current genetic research. Another unsolved question in genetics is whether or not
epigenetics is important in evolution. Epigenetics is when a trait is inherited without there being any change
in gene sequences.[37]

[edit] Variation
Further information: Genetic diversity and Population genetics
An individual organism's phenotype results from both its genotype and the influence from the
environment it has lived in. A substantial part of the variation in phenotypes in a population is caused by the
differences between their genotypes.[36] The modern evolutionary synthesis defines evolution as the change
over time in this genetic variation. The frequency of one particular allele will fluctuate, becoming more or less
prevalent relative to other forms of that gene. Evolutionary forces act by driving these changes in allele
frequency in one direction or another. Variation disappears when a new allele reaches the point of fixation —
when it either disappears from the population or replaces the ancestral allele entirely.[38]
Variation comes from mutations in genetic material, migration between populations (gene flow), and
the reshuffling of genes through sexual reproduction. Variation also comes from exchanges of genes
between different species; for example, through horizontal gene transfer in bacteria, and hybridization in
plants.[39] Despite the constant introduction of variation through these processes, most of the genome of a
species is identical in all individuals of that species.[40] However, even relatively small changes in genotype
can lead to dramatic changes in phenotype: for example, chimpanzees and humans differ in only about 5%
of their genomes.[41]
[edit] Mutation
Further information: Mutation and Molecular evolution

Duplication of part of a chromosome

Random mutations constantly occur in the genomes of organisms; these mutations create genetic
variation. Mutations are changes in the DNA sequence of a cell's genome and are caused by radiation,
viruses, transposons and mutagenic chemicals, as well as errors that occur during meiosis or DNA
replication.[42][43][44] These mutations involve several different types of change in DNA sequences; these
can either have no effect, alter the product of a gene, or prevent the gene from functioning. Studies in the fly
Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene, this will probably
be harmful, with about 70 percent of these mutations having damaging effects, and the remainder being
either neutral or weakly beneficial.[45] Due to the damaging effects that mutations can have on cells,
organisms have evolved mechanisms such as DNA repair to remove mutations.[42] Therefore, the optimal
mutation rate for a species is a trade-off between costs of a high mutation rate, such as deleterious
mutations, and the metabolic costs of maintaining systems to reduce the mutation rate, such as DNA repair
enzymes.[46] Viruses that use RNA as their genetic material have rapid mutation rates,[47] which can be an
advantage since these viruses will evolve constantly and rapidly, and thus evade the defensive responses of
e.g. the human immune system.[48]
Mutations can involve large sections of a chromosome becoming duplicated (usually by genetic
recombination), which can introduce extra copies of a gene into a genome.[49] Extra copies of genes are a
major source of the raw material needed for new genes to evolve.[50] This is important because most new
genes evolve within gene families from pre-existing genes that share common ancestors.[51] For example,
the human eye uses four genes to make structures that sense light: three for color vision and one for night
vision; all four are descended from a single ancestral gene.[52] New genes can be created from an ancestral
gene when a duplicate copy mutates and acquires a new function. This process is easier once a gene has
been duplicated because it increases the redundancy of the system; one gene in the pair can acquire a new
function while the other copy continues to perform its original function.[53][54] Other types of mutation can
even create entirely new genes from previously noncoding DNA.[55][56] The creation of new genes can also
involve small parts of several genes being duplicated, with these fragments then recombining to form new
combinations with new functions.[57][58] When new genes are assembled from shuffling pre-existing parts,
domains act as modules with simple independent functions, which can be mixed together creating new
combinations with new and complex functions.[59] For example, polyketide synthases are large enzymes
that make antibiotics; they contain up to one hundred independent domains that each catalyze one step in
the overall process, like a step in an assembly line.[60]
Changes in chromosome number may involve even larger mutations, where segments of the DNA
within chromosomes break and then rearrange. For example, two chromosomes in the Homo genus fused to
produce human chromosome 2; this fusion did not occur in the lineage of the other apes, and they retain
these separate chromosomes.[61] In evolution, the most important role of such chromosomal
rearrangements may be to accelerate the divergence of a population into new species by making populations
less likely to interbreed, and thereby preserving genetic differences between these populations.[62]
Sequences of DNA that can move about the genome, such as transposons, make up a major fraction
of the genetic material of plants and animals, and may have been important in the evolution of genomes.[63]
For example, more than a million copies of the Alu sequence are present in the human genome, and these
sequences have now been recruited to perform functions such as regulating gene expression.[64] Another
effect of these mobile DNA sequences is that when they move within a genome, they can mutate or delete
existing genes and thereby produce genetic diversity.[43]

[edit] Sex and recombination

Further information: Genetic recombination and Sexual reproduction
In asexual organisms, genes are inherited together, or linked, as they cannot mix with genes of other
organisms during reproduction. In contrast, the offspring of sexual organisms contain random mixtures of
their parents' chromosomes that are produced through independent assortment. In a related process called
homologous recombination, sexual organisms exchange DNA between two matching chromosomes.[65]
Recombination and reassortment do not alter allele frequencies, but instead change which alleles are
associated with each other, producing offspring with new combinations of alleles.[66] Sex usually increases
genetic variation and may increase the rate of evolution.[67][68] However, asexuality is advantageous in
some environments as it can evolve in previously sexual animals.[69] Here, asexuality might allow the two
sets of alleles in their genome to diverge and gain different functions.[70]
Recombination allows even alleles that are close together in a strand of DNA to be inherited
independently. However, the rate of recombination is low (approximately two events per chromosome per
generation). As a result, genes close together on a chromosome may not always be shuffled away from each
other, and genes that are close together tend to be inherited together, a phenomenon known as linkage.[71]
This tendency is measured by finding how often two alleles occur together on a single chromosome, which is
called their linkage disequilibrium. A set of alleles that is usually inherited in a group is called a haplotype.
This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can
drive a selective sweep that will also cause the other alleles in the haplotype to become more common in the
population; this effect is called genetic hitchhiking.[72]
When alleles cannot be separated by recombination – such as in mammalian Y chromosomes, which
pass intact from fathers to sons – harmful mutations accumulate.[73][74] By breaking up allele combinations,
sexual reproduction allows the removal of harmful mutations and the retention of beneficial mutations.[75] In
addition, recombination and reassortment can produce individuals with new and advantageous gene
combinations. These positive effects are balanced by the fact that sex reduces an organism's reproductive
rate, can cause mutations and may separate beneficial combinations of genes.[75] The reasons for the
evolution of sexual reproduction are therefore unclear and this question is still an active area of research in
evolutionary biology,[76][77] that has prompted ideas such as the Red Queen hypothesis.[78]

[edit] Population genetics

White peppered moth

Black morph in peppered moth evolution
Further information: Population genetics
From a genetic viewpoint, evolution is a generation-to-generation change in the frequencies of
alleles within a population that shares a common gene pool .[79] A population is a localized group of
individuals belonging to the same species. For example, all of the moths of the same species living in an
isolated forest represent a population. A single gene in this population may have several alternate forms,
which account for variations between the phenotypes of the organisms. An example might be a gene for
coloration in moths that has two alleles: black and white. A gene pool is the complete set of alleles for a gene
in a single population; the allele frequency measures the fraction of the gene pool composed of a single allele
(for example, what fraction of moth coloration genes are the black allele). Evolution occurs when there are
changes in the frequencies of alleles within a population of interbreeding organisms; for example, the allele
for black color in a population of moths becoming more common.
To understand the mechanisms that cause a population to evolve, it is useful to consider what
conditions are required for a population not to evolve. The Hardy-Weinberg principle states that the
frequencies of alleles (variations in a gene) in a sufficiently large population will remain constant if the only
forces acting on that population are the random reshuffling of alleles during the formation of the sperm or
egg, and the random combination of the alleles in these sex cells during fertilization.[80] Such a population is
said to be in Hardy-Weinberg equilibrium; it is not evolving.[81]

[edit] Gene flow

Further information: Gene flow, Hybrid (biology), and Horizontal gene transfer
When they mature, male lions leave the pride where they were born and take over a new pride to
mate, causing gene flow between prides.[82]
Gene flow is the exchange of genes between populations, which are usually of the same species.[83]
Examples of gene flow within a species include the migration and then breeding of organisms, or the
exchange of pollen. Gene transfer between species includes the formation of hybrid organisms and
horizontal gene transfer.
Migration into or out of a population can change allele frequencies, as well as introducing genetic
variation into a population. Immigration may add new genetic material to the established gene pool of a
population. Conversely, emigration may remove genetic material. As barriers to reproduction between two
diverging populations are required for the populations to become new species, gene flow may slow this
process by spreading genetic differences between the populations. Gene flow is hindered by mountain
ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which has
hindered the flow of plant genes.[84]
Depending on how far two species have diverged since their most recent common ancestor, it may
still be possible for them to produce offspring, as with horses and donkeys mating to produce mules.[85]
Such hybrids are generally infertile, due to the two different sets of chromosomes being unable to pair up
during meiosis. In this case, closely related species may regularly interbreed, but hybrids will be selected
against and the species will remain distinct. However, viable hybrids are occasionally formed and these new
species can either have properties intermediate between their parent species, or possess a totally new
phenotype.[86] The importance of hybridization in creating new species of animals is unclear, although cases
have been seen in many types of animals,[87] with the gray tree frog being a particularly well-studied
Hybridization is, however, an important means of speciation in plants, since polyploidy (having more
than two copies of each chromosome) is tolerated in plants more readily than in animals.[89][90] Polyploidy is
important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to
pair with an identical partner during meiosis.[91] Polyploids also have more genetic diversity, which allows
them to avoid inbreeding depression in small populations.[92]
Horizontal gene transfer is the transfer of genetic material from one organism to another organism
that is not its offspring; this is most common among bacteria.[93] In medicine, this contributes to the spread
of antibiotic resistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other
species.[2] Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces
cerevisiae and the adzuki bean beetle Callosobruchus chinensis may also have occurred.[94][95] An
example of larger-scale transfers are the eukaryotic bdelloid rotifers, which appear to have received a range
of genes from bacteria, fungi, and plants.[96] Viruses can also carry DNA between organisms, allowing
transfer of genes even across biological domains.[97] Large-scale gene transfer has also occurred between
the ancestors of eukaryotic cells and prokaryotes, during the acquisition of chloroplasts and mitochondria.

[edit] Mechanisms
The two main mechanisms that produce evolution are natural selection and genetic drift. Natural
selection is the process which favors genes that aid survival and reproduction. Genetic drift is the random
change in the frequency of alleles, caused by the random sampling of a generation's genes during
reproduction. The relative importance of natural selection and genetic drift in a population varies depending
on the strength of the selection and the effective population size, which is the number of individuals capable
of breeding.[99] Natural selection usually predominates in large populations, whereas genetic drift dominates
in small populations. The dominance of genetic drift in small populations can even lead to the fixation of
slightly deleterious mutations.[100] As a result, changing population size can dramatically influence the
course of evolution. Population bottlenecks, where the population shrinks temporarily and therefore loses
genetic variation, result in a more uniform population.[38]

[edit] Natural selection

Further information: Natural selection and Fitness (biology)
Natural selection of a population for dark coloration.
Natural selection is the process by which genetic mutations that enhance reproduction become, and
remain, more common in successive generations of a population. It has often been called a "self-evident"
mechanism because it necessarily follows from three simple facts:
• Heritable variation exists within populations of organisms.
• Organisms produce more offspring than can survive.
• These offspring vary in their ability to survive and reproduce.
These conditions produce competition between organisms for survival and reproduction.
Consequently, organisms with traits that give them an advantage over their competitors pass these
advantageous traits on, while traits that do not confer an advantage are not passed on to the next generation.
The central concept of natural selection is the evolutionary fitness of an organism.[102] Fitness is
measured by an organism's ability to survive and reproduce, which determines the size of its genetic
contribution to the next generation.[102] However, fitness is not the same as the total number of offspring:
instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes.[103]
For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and
weak to survive, this organism would make little genetic contribution to future generations and would thus
have low fitness.[102]
If an allele increases fitness more than the other alleles of that gene, then with each generation this
allele will become more common within the population. These traits are said to be "selected for". Examples of
traits that can increase fitness are enhanced survival, and increased fecundity. Conversely, the lower fitness
caused by having a less beneficial or deleterious allele results in this allele becoming rarer — they are
"selected against".[5] Importantly, the fitness of an allele is not a fixed characteristic; if the environment
changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become
harmful.[1] However, even if the direction of selection does reverse in this way, traits that were lost in the past
may not re-evolve in an identical form (see Dollo's law).[104][105]
A chart showing three types of selection. 1.Disruptive selection 2.Stabilizing selection 3.Directional
Natural selection within a population for a trait that can vary across a range of values, such as height,
can be categorized into three different types. The first is directional selection, which is a shift in the average
value of a trait over time — for example organisms slowly getting taller.[106] Secondly, disruptive selection is
selection for extreme trait values and often results in two different values becoming most common, with
selection against the average value. This would be when either short or tall organisms had an advantage, but
not those of medium height. Finally, in stabilizing selection there is selection against extreme trait values on
both ends, which causes a decrease in variance around the average value and less diversity.[101][107] This
would, for example, cause organisms to slowly become all the same height.
A special case of natural selection is sexual selection, which is selection for any trait that increases
mating success by increasing the attractiveness of an organism to potential mates.[108] Traits that evolved
through sexual selection are particularly prominent in males of some animal species, despite traits such as
cumbersome antlers, mating calls or bright colors that attract predators, decreasing the survival of individual
males.[109] This survival disadvantage is balanced by higher reproductive success in males that show these
hard to fake, sexually selected traits.[110]
Natural selection most generally makes nature the measure against which individuals, and individual
traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem, that is, a system in
which organisms interact with every other element, physical as well as biological, in their local environment.
Eugene Odum, a founder of ecology, defined an ecosystem as: "Any unit that includes all of the a given area interacting with the physical environment so that a flow of energy leads to clearly
defined trophic structure, biotic diversity, and material cycles (ie: exchange of materials between living and
nonliving parts) within the system."[111] Each population within an ecosystem occupies a distinct niche, or
position, with distinct relationships to other parts of the system. These relationships involve the life history of
the organism, its position in the food chain, and its geographic range. This broad understanding of nature
enables scientists to delineate specific forces which, together, comprise natural selection.
An active area of research is the unit of selection, with natural selection being proposed to work at
the level of genes, cells, individual organisms, groups of organisms and species.[112][113] None of these are
mutually exclusive and selection can act on multiple levels simultaneously.[114] An example of selection
occurring below the level of the individual organism are genes called transposons, which can replicate and
spread throughout a genome.[115] Selection at a level above the individual, such as group selection, may
allow the evolution of co-operation, as discussed below.[116]

[edit] Genetic drift

Further information: Genetic drift and Effective population size
Simulation of genetic drift of 20 unlinked alleles in populations of 10 (top) and 100 (bottom). Drift to
fixation is more rapid in the smaller population.
Genetic drift is the change in allele frequency from one generation to the next that occurs because
alleles in offspring are a random sample of those in the parents, as well as from the role that chance plays in
determining whether a given individual will survive and reproduce. In mathematical terms, alleles are subject
to sampling error. As a result, when selective forces are absent or relatively weak, allele frequencies tend to
"drift" upward or downward randomly (in a random walk). This drift halts when an allele eventually becomes
fixed, either by disappearing from the population, or replacing the other alleles entirely. Genetic drift may
therefore eliminate some alleles from a population due to chance alone. Even in the absence of selective
forces, genetic drift can cause two separate populations that began with the same genetic structure to drift
apart into two divergent populations with different sets of alleles.[117]
The time for an allele to become fixed by genetic drift depends on population size, with fixation
occurring more rapidly in smaller populations.[118] The precise measure of population that is important is
called the effective population size. The effective population is always smaller than the total population since
it takes into account factors such as the level of inbreeding, the number of animals that are too old or young
to breed, and the lower probability of animals that live far apart managing to mate with each other.[119]
An example when genetic drift is probably of central importance in determining a trait is the loss of
pigments from animals that live in caves, a change that produces no obvious advantage or disadvantage in
complete darkness.[120] However, it is usually difficult to measure the relative importance of selection and
drift,[121] so the comparative importance of these two forces in driving evolutionary change is an area of
current research.[122] These investigations were prompted by the neutral theory of molecular evolution,
which proposed that most evolutionary changes are the result of the fixation of neutral mutations that do not
have any immediate effects on the fitness of an organism.[123] Hence, in this model, most genetic changes
in a population are the result of constant mutation pressure and genetic drift.[124] This form of the neutral
theory is now largely abandoned, since it does not seem to fit the genetic variation seen in nature.[125][126]
However, a more recent and better-supported version of this model is the nearly neutral theory, where most
mutations only have small effects on fitness.[101]
[edit] Outcomes
Evolution influences every aspect of the form and behavior of organisms. Most prominent are the
specific behavioral and physical adaptations that are the outcome of natural selection. These adaptations
increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms
can also respond to selection by co-operating with each other, usually by aiding their relatives or engaging in
mutually beneficial symbiosis. In the longer term, evolution produces new species through splitting ancestral
populations of organisms into new groups that cannot or will not interbreed.
These outcomes of evolution are sometimes divided into macroevolution, which is evolution that
occurs at or above the level of species, such as extinction and speciation, and microevolution, which is
smaller evolutionary changes, such as adaptations, within a species or population.[127] In general,
macroevolution is regarded as the outcome of long periods of microevolution.[128] Thus, the distinction
between micro- and macroevolution is not a fundamental one – the difference is simply the time involved.
[129] However, in macroevolution, the traits of the entire species may be important. For instance, a large
amount of variation among individuals allows a species to rapidly adapt to new habitats, lessening the
chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it
more likely that part of the population will become isolated. In this sense, microevolution and macroevolution
might involve selection at different levels – with microevolution acting on genes and organisms, versus
macroevolutionary processes such as species selection acting on entire species and affecting their rates of
speciation and extinction.[130][131][132]
A common misconception is that evolution has goals or long-term plans; realistically however,
evolution has no long-term goal and does not necessarily produce greater complexity.[133][134] Although
complex species have evolved, they occur as a side effect of the overall number of organisms increasing,
and simple forms of life still remain more common in the biosphere.[135] For example, the overwhelming
majority of species are microscopic prokaryotes, which form about half the world's biomass despite their
small size,[136] and constitute the vast majority of Earth's biodiversity.[137] Simple organisms have therefore
been the dominant form of life on Earth throughout its history and continue to be the main form of life up to
the present day, with complex life only appearing more diverse because it is more noticeable.[138] Indeed,
the evolution of microorganisms is particularly important to modern evolutionary research, since their rapid
reproduction allows the study of experimental evolution and the observation of evolution and adaptation in
real time.[139][140]

[edit] Adaptation
For more details on this topic, see Adaptation.
Adaptation is one of the basic phenomena of biology,[141] and is the process whereby an organism
becomes better suited to its habitat.[142][143] Also, the term adaptation may refer to a trait that is important
for an organism's survival. For example, the adaptation of horses' teeth to the grinding of grass, or the ability
of horses to run fast and escape predators. By using the term adaptation for the evolutionary process, and
adaptive trait for the product (the bodily part or function), the two senses of the word may be distinguished.
Adaptations are produced by natural selection.[144] The following definitions are due to Theodosius
1. Adaptation is the evolutionary process whereby an organism becomes better able to live in
its habitat or habitats.[145]
2. Adaptedness is the state of being adapted: the degree to which an organism is able to live
and reproduce in a given set of habitats.[146]
3. An adaptive trait is an aspect of the developmental pattern of the organism which enables or
enhances the probability of that organism surviving and reproducing.[147]
Adaptation may cause either the gain of a new feature, or the loss of an ancestral feature. An
example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes
causing antibiotic resistance by both modifying the target of the drug, or increasing the activity of transporters
that pump the drug out of the cell.[148] Other striking examples are the bacteria Escherichia coli evolving the
ability to use citric acid as a nutrient in a long-term laboratory experiment,[149] Flavobacterium evolving a
novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing,[150][151] and
the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic
pesticide pentachlorophenol.[152][153] An interesting but still controversial idea is that some adaptations
might increase the ability of organisms to generate genetic diversity and adapt by natural selection
(increasing organisms' evolvability).[154][155]

A baleen whale skeleton, a and b label flipper bones, which were adapted from front leg bones: while
c indicates vestigial leg bones, suggesting an adaptation from land to sea.[156]
Adaptation occurs through the gradual modification of existing structures. Consequently, structures
with similar internal organization may have different functions in related organisms. This is the result of a
single ancestral structure being adapted to function in different ways. The bones within bat wings, for
example, are very similar to those in mice feet and primate hands, due to the descent of all these structures
from a common mammalian ancestor.[157] However, since all living organisms are related to some extent,
[158] even organs that appear to have little or no structural similarity, such as arthropod, squid and vertebrate
eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous
genes that control their assembly and function; this is called deep homology.[159][160]
During adaptation, some structures may lose their original function and become vestigial structures.
[161] Such structures may have little or no function in a current species, yet have a clear function in ancestral
species, or other closely related species. Examples include pseudogenes,[162] the non-functional remains of
eyes in blind cave-dwelling fish,[163] wings in flightless birds,[164] and the presence of hip bones in whales
and snakes.[156] Examples of vestigial structures in humans include wisdom teeth,[165] the coccyx,[161] the
vermiform appendix,[161] and other behavioral vestiges such as goose bumps,[166] and primitive reflexes.
However, many traits that appear to be simple adaptations are in fact exaptations: structures
originally adapted for one function, but which coincidentally became somewhat useful for some other function
in the process.[171] One example is the African lizard Holaspis guentheri, which developed an extremely flat
head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head
has become so flattened that it assists in gliding from tree to tree—an exaptation.[171] Within cells, molecular
machines such as the bacterial flagella[172] and protein sorting machinery[173] evolved by the recruitment of
several pre-existing proteins that previously had different functions.[127] Another example is the recruitment
of enzymes from glycolysis and xenobiotic metabolism to serve as structural proteins called crystallins within
the lenses of organisms' eyes.[174][175]
A critical principle of ecology is that of competitive exclusion: no two species can occupy the same
niche in the same environment for a long time.[176] Consequently, natural selection will tend to force species
to adapt to different ecological niches. This may mean that, for example, two species of cichlid fish adapt to
live in different habitats, which will minimize the competition between them for food.[177]
An area of current investigation in evolutionary developmental biology is the developmental basis of
adaptations and exaptations.[178] This research addresses the origin and evolution of embryonic
development and how modifications of development and developmental processes produce novel features.
[179] These studies have shown that evolution can alter development to create new structures, such as
embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in
mammals.[180] It is also possible for structures that have been lost in evolution to reappear due to changes
in developmental genes, such as a mutation in chickens causing embryos to grow teeth similar to those of
crocodiles.[181] It is now becoming clear that most alterations in the form of organisms are due to changes in
a small set of conserved genes.[182]
[edit] Co-evolution

Common garter snake (Thamnophis sirtalis sirtalis) which has evolved resistance to tetrodotoxin in
its amphibian prey.
Further information: Co-evolution
Interactions between organisms can produce both conflict and co-operation. When the interaction is
between pairs of species, such as a pathogen and a host, or a predator and its prey, these species can
develop matched sets of adaptations. Here, the evolution of one species causes adaptations in a second
species. These changes in the second species then, in turn, cause new adaptations in the first species. This
cycle of selection and response is called co-evolution.[183] An example is the production of tetrodotoxin in
the rough-skinned newt and the evolution of tetrodotoxin resistance in its predator, the common garter snake.
In this predator-prey pair, an evolutionary arms race has produced high levels of toxin in the newt and
correspondingly high levels of toxin resistance in the snake.[184]

[edit] Co-operation
Further information: Co-operation (evolution)
However, not all interactions between species involve conflict.[185] Many cases of mutually
beneficial interactions have evolved. For instance, an extreme cooperation exists between plants and the
mycorrhizal fungi that grow on their roots and aid the plant in absorbing nutrients from the soil.[186] This is a
reciprocal relationship as the plants provide the fungi with sugars from photosynthesis. Here, the fungi
actually grow inside plant cells, allowing them to exchange nutrients with their hosts, while sending signals
that suppress the plant immune system.[187]
Coalitions between organisms of the same species have also evolved. An extreme case is the
eusociality found in social insects, such as bees, termites and ants, where sterile insects feed and guard the
small number of organisms in a colony that are able to reproduce. On an even smaller scale, the somatic
cells that make up the body of an animal limit their reproduction so they can maintain a stable organism,
which then supports a small number of the animal's germ cells to produce offspring. Here, somatic cells
respond to specific signals that instruct them whether to grow, remain as they are, or die. If cells ignore these
signals and multiply inappropriately, their uncontrolled growth causes cancer.[42]
Such cooperation within species may have evolved through the process of kin selection, which is
where one organism acts to help raise a relative's offspring.[188] This activity is selected for because if the
helping individual contains alleles which promote the helping activity, it is likely that its kin will also contain
these alleles and thus those alleles will be passed on.[189] Other processes that may promote cooperation
include group selection, where cooperation provides benefits to a group of organisms.[190]

[edit] Speciation
Further information: Speciation
The four mechanisms of speciation.
Speciation is the process where a species diverges into two or more descendant species.[191]
Evolutionary biologists view species as statistical phenomena and not categories or types. This view is
counterintuitive since the classical idea of species is still widely held, with a species seen as a class of
organisms exemplified by a "type specimen" that bears all the traits common to this species. Instead, a
species is now defined as a separately evolving lineage that forms a single gene pool. Although properties
such as genetics and morphology are used to help separate closely related lineages, this definition has fuzzy
boundaries.[192] Indeed, the exact definition of the term "species" is still controversial, particularly in
prokaryotes,[193] and this is called the species problem.[194] Biologists have proposed a range of more
precise definitions, but the definition used is a pragmatic choice that depends on the particularities of the
species concerned.[194] Typically the actual focus on biological study is the population, an observable
interacting group of organisms, rather than a species, an observable similar group of individuals.
Speciation has been observed multiple times under both controlled laboratory conditions and in
nature.[195] In sexually reproducing organisms, speciation results from reproductive isolation followed by
genealogical divergence. There are four mechanisms for speciation. The most common in animals is
allopatric speciation, which occurs in populations initially isolated geographically, such as by habitat
fragmentation or migration. Selection under these conditions can produce very rapid changes in the
appearance and behaviour of organisms.[196][197] As selection and drift act independently on populations
isolated from the rest of their species, separation may eventually produce organisms that cannot interbreed.
The second mechanism of speciation is peripatric speciation, which occurs when small populations
of organisms become isolated in a new environment. This differs from allopatric speciation in that the isolated
populations are numerically much smaller than the parental population. Here, the founder effect causes rapid
speciation through both rapid genetic drift and selection on a small gene pool.[199]
The third mechanism of speciation is parapatric speciation. This is similar to peripatric speciation in
that a small population enters a new habitat, but differs in that there is no physical separation between these
two populations. Instead, speciation results from the evolution of mechanisms that reduce gene flow between
the two populations.[191] Generally this occurs when there has been a drastic change in the environment
within the parental species' habitat. One example is the grass Anthoxanthum odoratum, which can undergo
parapatric speciation in response to localized metal pollution from mines.[200] Here, plants evolve that have
resistance to high levels of metals in the soil. Selection against interbreeding with the metal-sensitive
parental population produced a gradual change in the flowering time of the metal-resistant plants, which
eventually produced complete reproductive isolation. Selection against hybrids between the two populations
may cause reinforcement, which is the evolution of traits that promote mating within a species, as well as
character displacement, which is when two species become more distinct in appearance.[201]
Geographical isolation of finches on the Galápagos Islands produced over a dozen new species.
Finally, in sympatric speciation species diverge without geographic isolation or changes in habitat.
This form is rare since even a small amount of gene flow may remove genetic differences between parts of a
population.[202] Generally, sympatric speciation in animals requires the evolution of both genetic differences
and non-random mating, to allow reproductive isolation to evolve.[203]
One type of sympatric speciation involves cross-breeding of two related species to produce a new
hybrid species. This is not common in animals as animal hybrids are usually sterile. This is because during
meiosis the homologous chromosomes from each parent are from different species and cannot successfully
pair. However, it is more common in plants because plants often double their number of chromosomes, to
form polyploids.[204] This allows the chromosomes from each parental species to form matching pairs during
meiosis, since each parent's chromosomes are represented by a pair already.[205] An example of such a
speciation event is when the plant species Arabidopsis thaliana and Arabidopsis arenosa cross-bred to give
the new species Arabidopsis suecica.[206] This happened about 20,000 years ago,[207] and the speciation
process has been repeated in the laboratory, which allows the study of the genetic mechanisms involved in
this process.[208] Indeed, chromosome doubling within a species may be a common cause of reproductive
isolation, as half the doubled chromosomes will be unmatched when breeding with undoubled organisms.[90]
Speciation events are important in the theory of punctuated equilibrium, which accounts for the
pattern in the fossil record of short "bursts" of evolution interspersed with relatively long periods of stasis,
where species remain relatively unchanged.[209] In this theory, speciation and rapid evolution are linked,
with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel
habitats or small populations. As a result, the periods of stasis in the fossil record correspond to the parental
population, and the organisms undergoing speciation and rapid evolution are found in small populations or
geographically restricted habitats, and therefore rarely being preserved as fossils.[210]

[edit] Extinction
Further information: Extinction
Tyrannosaurus rex. Non-avian dinosaurs died out in the Cretaceous–Tertiary extinction event at the
end of the Cretaceous period.
Extinction is the disappearance of an entire species. Extinction is not an unusual event, as species
regularly appear through speciation, and disappear through extinction.[211] Nearly all animal and plant
species that have lived on earth are now extinct,[212] and extinction appears to be the ultimate fate of all
species.[213] These extinctions have happened continuously throughout the history of life, although the rate
of extinction spikes in occasional mass extinction events.[214] The Cretaceous–Tertiary extinction event,
during which the non-avian dinosaurs went extinct, is the most well-known, but the earlier Permian–Triassic
extinction event was even more severe, with approximately 96 percent of species driven to extinction.[214]
The Holocene extinction event is an ongoing mass extinction associated with humanity's expansion across
the globe over the past few thousand years. Present-day extinction rates are 100–1000 times greater than the
background rate, and up to 30 percent of species may be extinct by the mid 21st century.[215] Human
activities are now the primary cause of the ongoing extinction event;[216] global warming may further
accelerate it in the future.[217]
The role of extinction in evolution is not very well understood and may depend on which type of
extinction is considered.[214] The causes of the continuous "low-level" extinction events, which form the
majority of extinctions, may be the result of competition between species for limited resources (competitive
exclusion).[14] If one species can out-compete another, this could produce species selection, with the fitter
species surviving and the other species being driven to extinction.[112] The intermittent mass extinctions are
also important, but instead of acting as a selective force, they drastically reduce diversity in a nonspecific
manner and promote bursts of rapid evolution and speciation in survivors.[218]

[edit] Evolutionary history of life

Main article: Evolutionary history of life
See also: Timeline of evolution and Timeline of human evolution

[edit] Origin of life

Further information: Abiogenesis and RNA world hypothesis
The origin of life is a necessary precursor for biological evolution, but understanding that evolution
occurred once organisms appeared and investigating how this happens does not depend on understanding
exactly how life began.[219] The current scientific consensus is that the complex biochemistry that makes up
life came from simpler chemical reactions, but it is unclear how this occurred.[220] Not much is certain about
the earliest developments in life, the structure of the first living things, or the identity and nature of any last
universal common ancestor or ancestral gene pool.[221][222] Consequently, there is no scientific consensus
on how life began, but proposals include self-replicating molecules such as RNA,[223] and the assembly of
simple cells.[224]

[edit] Common descent

Further information: Common descent, Evidence of common descent, and Homology (biology)

The hominoids are descendants of a common ancestor.

All organisms on Earth are descended from a common ancestor or ancestral gene pool.[158][225]
Current species are a stage in the process of evolution, with their diversity the product of a long series of
speciation and extinction events.[226] The common descent of organisms was first deduced from four simple
facts about organisms: First, they have geographic distributions that cannot be explained by local adaptation.
Second, the diversity of life is not a set of completely unique organisms, but organisms that share
morphological similarities. Third, vestigial traits with no clear purpose resemble functional ancestral traits,
and finally, that organisms can be classified using these similarities into a hierarchy of nested groups – similar
to a family tree.[9] However, modern research has suggested that, due to horizontal gene transfer, this "tree
of life" may be more complicated than a simple branching tree since some genes have spread independently
between distantly related species.[227][228]
Past species have also left records of their evolutionary history. Fossils, along with the comparative
anatomy of present-day organisms, constitute the morphological, or anatomical, record.[229] By comparing
the anatomies of both modern and extinct species, paleontologists can infer the lineages of those species.
However, this approach is most successful for organisms that had hard body parts, such as shells, bones or
teeth. Further, as prokaryotes such as bacteria and archaea share a limited set of common morphologies,
their fossils do not provide information on their ancestry.
More recently, evidence for common descent has come from the study of biochemical similarities
between organisms. For example, all living cells use the same basic set of nucleotides and amino acids.[230]
The development of molecular genetics has revealed the record of evolution left in organisms' genomes:
dating when species diverged through the molecular clock produced by mutations.[231] For example, these
DNA sequence comparisons have revealed that humans and chimpanzees share 96% of their genomes and
analyzing the few areas where they differ helps shed light on when the common ancestor of these species

[edit] Evolution of life

For more details on this topic, see Timeline of evolution.
Evolutionary tree showing the divergence of modern species from their common ancestor in the
center.[233] The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.
Despite the uncertainty on how life began, it is generally accepted that prokaryotes inhabited the
Earth from approximately 3–4 billion years ago.[234][235] No obvious changes in morphology or cellular
organization occurred in these organisms over the next few billion years.[236]
The eukaryotes were the next major change in cell structure. These came from ancient bacteria
being engulfed by the ancestors of eukaryotic cells, in a cooperative association called endosymbiosis.[98]
[237] The engulfed bacteria and the host cell then underwent co-evolution, with the bacteria evolving into
either mitochondria or hydrogenosomes.[238] An independent second engulfment of cyanobacterial-like
organisms led to the formation of chloroplasts in algae and plants.[239] It is unknown when the first
eukaryotic cells appeared though they first emerged between 1.6 – 2.7 billion years ago.
The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until about 610
million years ago when multicellular organisms began to appear in the oceans in the Ediacaran period.[234]
[240] The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as
sponges, brown algae, cyanobacteria, slime moulds and myxobacteria.[241]
Soon after the emergence of these first multicellular organisms, a remarkable amount of biological
diversity appeared over approximately 10 million years, in an event called the Cambrian explosion. Here, the
majority of types of modern animals appeared in the fossil record, as well as unique lineages that
subsequently became extinct.[242] Various triggers for the Cambrian explosion have been proposed,
including the accumulation of oxygen in the atmosphere from photosynthesis.[243] About 500 million years
ago, plants and fungi colonized the land, and were soon followed by arthropods and other animals.[244]
Insects were particularly successful and even today make up the majority of animal species.[245]
Amphibians first appeared around 300 million years ago, followed by early amniotes, then mammals around
200 million years ago and birds around 100 million years ago (both from "reptile"-like lineages). However,
despite the evolution of these large animals, smaller organisms similar to the types that evolved early in this
process continue to be highly successful and dominate the Earth, with the majority of both biomass and
species being prokaryotes.[137]
[edit] Applications
Further information: Artificial selection and Evolutionary computation
Evolutionary biology, and in particular the understanding of how organisms evolve through natural
selection, is an area of science with many practical applications.[246] A major technological application of
evolution is artificial selection, which is the intentional selection of certain traits in a population of organisms.
Humans have used artificial selection for thousands of years in the domestication of plants and animals.[247]
More recently, such selection has become a vital part of genetic engineering, with selectable markers such
as antibiotic resistance genes being used to manipulate DNA in molecular biology. It is also possible to use
repeated rounds of mutation and selection to evolve proteins with particular properties, such as modified
enzymes or new antibodies, in a process called directed evolution.[248]
Understanding the changes that have occurred during organism's evolution can reveal the genes
needed to construct parts of the body, genes which may be involved in human genetic disorders.[249] For
example, the Mexican tetra is an albino cavefish that lost its eyesight during evolution. Breeding together
different populations of this blind fish produced some offspring with functional eyes, since different mutations
had occurred in the isolated populations that had evolved in different caves.[250] This helped identify genes
required for vision and pigmentation, such as crystallins and the melanocortin 1 receptor.[251] Similarly,
comparing the genome of the Antarctic icefish, which lacks red blood cells, to close relatives such as the
zebrafish revealed genes needed to make these blood cells.[252]
As evolution can produce highly optimized processes and networks, it has many applications in
computer science. Here, simulations of evolution using evolutionary algorithms and artificial life started with
the work of Nils Aall Barricelli in the 1960s, and was extended by Alex Fraser, who published a series of
papers on simulation of artificial selection.[253] Artificial evolution became a widely recognized optimization
method as a result of the work of Ingo Rechenberg in the 1960s and early 1970s, who used evolution
strategies to solve complex engineering problems.[254] Genetic algorithms in particular became popular
through the writing of John Holland.[255] As academic interest grew, dramatic increases in the power of
computers allowed practical applications, including the automatic evolution of computer programs.[256]
Evolutionary algorithms are now used to solve multi-dimensional problems more efficiently than software
produced by human designers, and also to optimize the design of systems.[257]

[edit] Social and cultural responses

Further information: Social effect of evolutionary theory and Objections to evolution
As Darwinism became widely accepted in the 1870s, caricatures of Charles Darwin with an ape or
monkey body symbolised evolution.[258]
In the 19th century, particularly after the publication of On the Origin of Species in 1859, the idea that
life had evolved was an active source of academic debate centered on the philosophical, social and religious
implications of evolution. Nowadays, the fact that organisms evolve is uncontested in the scientific literature
and the modern evolutionary synthesis is widely accepted by scientists.[14] However, evolution remains a
contentious concept for some theists.[259]
While various religions and denominations have reconciled their beliefs with evolution through
concepts such as theistic evolution, there are creationists who believe that evolution is contradicted by the
creation myths found in their respective religions and who raise various objections to evolution.[127][260]
[261] As had been demonstrated by responses to the publication of Vestiges of the Natural History of
Creation in 1844, the most controversial aspect of evolutionary biology is the implication of human evolution
that human mental and moral faculties, which had been thought purely spiritual, are not distinctly separated
from those of other animals.[8] In some countries—notably the United States—these tensions between science
and religion have fueled the current creation-evolution controversy, a religious conflict focusing on politics
and public education.[262] While other scientific fields such as cosmology[263] and earth science[264] also
conflict with literal interpretations of many religious texts, evolutionary biology experiences significantly more
opposition from religious literalists.
The teaching of evolution in American secondary school biology classes was uncommon in most of
the first half of the 20th century. The Scopes Trial decision of 1925 caused the subject to become very rare in
American secondary biology textbooks for a generation, but it was gradually re-introduced about a generation
later and legally protected with the 1968 Epperson v. Arkansas decision. Since then, the competing religious
belief of creationism was legally disallowed in secondary school curricula in various decisions in the 1970s
and 1980s, but it returned in the form of intelligent design, to be excluded once again in the 2005 Kitzmiller v.
Dover Area School District case.[265]
Another example somewhat associated with evolutionary theory that is now widely regarded as
unwarranted is "Social Darwinism", a derogatory term associated with the 19th century Malthusian theory
developed by Whig philosopher Herbert Spencer. It was later expanded by others into ideas about "survival
of the fittest" in commerce and human societies as a whole, and led to claims that social inequality, sexism,
racism, and imperialism were justified.[266] However, these ideas contradict Darwin's own views, and
contemporary scientists and philosophers consider these ideas to be neither mandated by evolutionary
theory nor supported by data.[267][268][269]

[edit] See also


Books are collections of articles that can be downloaded or ordered in print.

[edit] References
1. ^ a b c Futuyma, Douglas J. (2005). Evolution. Sunderland, Massachusetts: Sinauer
Associates, Inc. ISBN 0-87893-187-2.
2. ^ a b Walsh T (2006). "Combinatorial genetic evolution of multiresistance". Curr. Opin.
Microbiol. 9 (5): 476–82. doi:10.1016/j.mib.2006.08.009. PMID 16942901.
3. ^ Kottak, Conrad Phillip. Window on Humanity: A Concise Introduction to Anthropology. New
York: The McGraw–Hill Companies, Inc, 2005
4. ^
5. ^ a b Lande R, Arnold SJ (1983). "The measurement of selection on correlated characters".
Evolution 37 (6): 1210–26. doi:10.2307/2408842.
6. ^ Ayala FJ (2007). "Darwin's greatest discovery: design without designer". Proc. Natl. Acad.
Sci. U.S.A. 104 (Suppl 1): 8567–73. doi:10.1073/pnas.0701072104. PMID 17494753. PMC 1876431.
7. ^ Ian C. Johnston (1999). "History of Science: Early Modern Geology". Malaspina University-
College. Retrieved 2008-01-15.
8. ^ a b Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press.
ISBN 0-52023693-9.
9. ^ a b Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray. p. 1.
ISBN 0801413192.
itemID=F373&viewtype=text&pageseq=16. . Related earlier ideas were acknowledged in Darwin,
Charles (1861). On the Origin of Species (3rd ed.). London: John Murray. xiii. ISBN 0801413192.
10.^ AAAS Council (December 26, 1922). "AAAS Resolution: Present Scientific Status of the
Theory of Evolution". American Association for the Advancement of Science.
11.^ a b "IAP Statement on the Teaching of Evolution" (PDF). The Interacademy Panel on
International Issues. 2006.
%20statement.pdf. Retrieved 2007-04-25. Joint statement issued by the national science academies
of 67 countries, including the United Kingdom's Royal Society
12.^ a b Board of Directors, American Association for the Advancement of Science (2006-02-16).
"Statement on the Teaching of Evolution" (PDF). American Association for the Advancement of
Science. from the world's
largest general scientific society
13.^ "Statements from Scientific and Scholarly Organizations". National Center for Science
14.^ a b c d e Kutschera U, Niklas K (2004). "The modern theory of biological evolution: an
expanded synthesis". Naturwissenschaften 91 (6): 255–76. doi:10.1007/s00114-004-0515-y.
PMID 15241603.
15.^ "Special report on evolution". New Scientist. 2008-01-19.
16.^ Wright, S (1984). Evolution and the Genetics of Populations, Volume 1: Genetic and
Biometric Foundations. The University of Chicago Press. ISBN 0-226-91038-5.
17.^ Zirkle C (1941). "Natural Selection before the "Origin of Species"". Proceedings of the
American Philosophical Society 84 (1): 71–123.
18.^ Muhammad Hamidullah and Afzal Iqbal (1993), The Emergence of Islam: Lectures on the
Development of Islamic World-view, Intellectual Tradition and Polity , p. 143-144. Islamic Research
Institute, Islamabad.
19.^ "A Source Book In Chinese Philosophy", Chan, Wing-Tsit, p. 204, 1962.
20.^ Terrall, M (2002). The Man Who Flattened the Earth: Maupertuis and the Sciences in the
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Genet. 5 (1): e1000326. doi:10.1371/journal.pgen.1000326. PMID 19119422.
252.^ Yergeau DA, Cornell CN, Parker SK, Zhou Y, Detrich HW (July 2005). "bloodthirsty, an
RBCC/TRIM gene required for erythropoiesis in zebrafish". Dev. Biol. 283 (1): 97–112.
doi:10.1016/j.ydbio.2005.04.006. PMID 15890331.
253.^ Fraser AS (1958). "Monte Carlo analyses of genetic models". Nature 181 (4603): 208–9.
doi:10.1038/181208a0. PMID 13504138.
254.^ Rechenberg, Ingo (1973) (in German). Evolutionsstrategie – Optimierung technischer
Systeme nach Prinzipien der biologischen Evolution (PhD thesis) . Fromman-Holzboog.
255.^ Holland, John H. (1975). Adaptation in Natural and Artificial Systems. University of
Michigan Press. ISBN 0262581116.
256.^ Koza, John R. (1992). Genetic Programming. MIT Press. ISBN 0262111705.
257.^ Jamshidi M (2003). "Tools for intelligent control: fuzzy controllers, neural networks and
genetic algorithms". Philosophical transactions. Series A, Mathematical, physical, and engineering
sciences 361 (1809): 1781–808. doi:10.1098/rsta.2003.1225. PMID 12952685.
258.^ Browne, Janet (2003). Charles Darwin: The Power of Place. London: Pimlico. pp. 376–
379. ISBN 0-7126-6837-3.
ISBN 978-0684824710.
*For the scientific and social reception of evolution in the 19th and early 20th centuries, see:
Johnston, Ian C.. "History of Science: Origins of Evolutionary Theory". And Still We Evolve. Liberal
Studies Department, Malaspina University College.
Retrieved 2007-05-24.
*Bowler, PJ (2003). Evolution: The History of an Idea, Third Edition, Completely Revised and
Expanded. University of California Press. ISBN 978-0520236936.
*Zuckerkandl E (2006). "Intelligent design and biological complexity". Gene 385: 2–18.
doi:10.1016/j.gene.2006.03.025. PMID 17011142.
260.^ Ross, M.R. (2005). "Who Believes What? Clearing up Confusion over Intelligent Design
and Young-Earth Creationism" (PDF). Journal of Geoscience Education 53 (3): 319. Retrieved 2008-04-28.
261.^ Hameed, Salman (2008-12-12). "Science and Religion: Bracing for Islamic Creationism".
Science 322 (5908): 1637–1638. doi:10.1126/science.1163672. PMID 19074331. Retrieved 2009.
262.^ Spergel D. N.; Scott, EC; Okamoto, S (2006). "Science communication. Public acceptance
of evolution". Science 313 (5788): 765–66. doi:10.1126/science.1126746. PMID 16902112.
263.^ Spergel, D. N.; Verde, L.; Peiris, H. V.; Komatsu, E.; Nolta, M. R.; Bennett, C. L.; Halpern,
M.; Hinshaw, G. et al. (2003). "First-Year Wilkinson Microwave Anisotropy Probe (WMAP)
Observations: Determination of Cosmological Parameters". The Astrophysical Journal Supplement
Series 148: 175–94. doi:10.1086/377226.
264.^ Wilde SA, Valley JW, Peck WH, Graham CM (2001). "Evidence from detrital zircons for
the existence of continental crust and oceans on the Earth 4.4 Gyr ago". Nature 409 (6817): 175–78.
doi:10.1038/35051550. PMID 11196637.
265.^ Understanding Creationism after Kitzmiller 2007
266.^ On the history of eugenics and evolution, see Kevles, D (1998). In the Name of Eugenics:
Genetics and the Uses of Human Heredity. Harvard University Press. ISBN 978-0674445574.
267.^ Darwin strongly disagreed with attempts by Herbert Spencer and others to extrapolate
evolutionary ideas to all possible subjects; see Midgley, M (2004). The Myths we Live By. Routledge.
p. 62. ISBN 978-0415340779.
268.^ Allhoff F (2003). "Evolutionary ethics from Darwin to Moore". History and philosophy of the
life sciences 25 (1): 51–79. doi:10.1080/03919710312331272945. PMID 15293515.
269.^ Gowaty, Patricia Adair (1997). Feminism and evolutionary biology: boundaries,
intersections, and frontiers. London: Chapman & Hall. ISBN 0-412-07361-7.

[edit] Further reading

Introductory reading
• Carroll, S. (2005). Endless Forms Most Beautiful. New York: W.W. Norton. ISBN 0-393-
• Charlesworth, C.B. and Charlesworth, D. (2003). Evolution. Oxfordshire: Oxford University
Press. ISBN 0-192-80251-8.
• Dawkins, R. (2006). The Selfish Gene: 30th Anniversary Edition. Oxford University Press.
ISBN 0199291152.
• Gould, S.J. (1989). Wonderful Life: The Burgess Shale and the Nature of History . New York:
W.W. Norton. ISBN 0-393-30700-X.
• Jones, S. (2001). Almost Like a Whale: The Origin of Species Updated. (American title:
Darwin's Ghost). New York: Ballantine Books. ISBN 0-345-42277-5.
• Mader, Sylvia S. (2007). Biology. Murray P. Pendarvis (9th ed.). McGraw Hill.
ISBN 9780073258393.
• Maynard Smith, J. (1993). The Theory of Evolution: Canto Edition. Cambridge University
Press. ISBN 0-521-45128-0.
• Pallen, M.J. (2009). The Rough Guide to Evolution. Rough Guides. ISBN 978-1-85828-946-
• Smith, C.B. and Sullivan, C. (2007). The Top 10 Myths about Evolution. Prometheus Books.
ISBN 978-1-59102-479-8.
History of evolutionary thought
• Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray.
ISBN 0801413192.
• Larson, E.J. (2004). Evolution: The Remarkable History of a Scientific Theory. New York:
Modern Library. ISBN 0-679-64288-9.
• Zimmer, C. (2001). Evolution: The Triumph of an Idea. London: HarperCollins. ISBN 0-060-
Advanced reading
• Barton, N.H., Briggs, D.E.G., Eisen, J.A., Goldstein, D.B. and Patel, N.H. (2007). Evolution.
Cold Spring Harbor Laboratory Press. ISBN 0-879-69684-2.
• Coyne, J.A. and Orr, H.A. (2004). Speciation. Sunderland: Sinauer Associates. ISBN 0-878-
• Futuyma, D.J. (2005). Evolution. Sunderland: Sinauer Associates. ISBN 0-878-93187-2.
• Gould, S.J. (2002). The Structure of Evolutionary Theory. Cambridge: Belknap Press
(Harvard University Press). ISBN 0-674-00613-5.
• Maynard Smith, J. and Szathmáry, E. (1997). The Major Transitions in Evolution.
Oxfordshire: Oxford University Press. ISBN 0-198-50294-X.
• Mayr, E. (2001). What Evolution Is. New York: Basic Books. ISBN 0-465-04426-3.
• Olson, Wendy; Hall, Brian Keith (2003). Keywords and concepts in evolutionary
developmental biology. Cambridge: Harvard University Press. ISBN 0-674-02240-8.

[edit] External links

General information
• Everything you wanted to know about evolution by New Scientist
• — How Evolution Works
• National Academies Evolution Resources
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and
• Understanding Evolution from University of California, Berkeley
• Evolution of Evolution - 150 Years of Darwin's "On the Origin of Species"
History of evolutionary thought
• The Complete Work of Charles Darwin Online
• Understanding Evolution: History, Theory, Evidence, and Implications
On-line lectures
• What Genomes Can Tell Us About the Past – lecture by Sydney Brenner
• The Origin of Vertebrates – lecture by Marc Kirschner
• The Making of the Fittest – lecture by Sean B. Carroll

Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

biology (Evo-devo) Canalisation · Modularity · Phenotypic plasticity
Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from ""

Categories: Articles with separate introductions | Evolution | Biology theories | Evolutionary biology

• This page was last modified on 21 August 2010 at 03:16.


Évolution (biologie)
Un article de Wikipédia, l'encyclopédie libre.

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Pour les articles homonymes, voir Évolution.

Arbre phylogénétique hypothétique de tous les organismes vivants. L'arbre est basé sur des
séquences de l'ARNr 16S. À l'origine proposé par Carl Woese, il montre l'histoire évolutive supposée des
trois domaines du vivant (bactéries, archées et eucaryotes).
En biologie, l'évolution désigne la transformation des espèces vivantes au cours des générations.
Cette transformation peut aboutir à la formation de nouvelles espèces, et donc à une diversification des
formes de vie. Cette diversification depuis les premières formes est à l'origine de la biodiversité actuelle.
L’histoire de l'évolution de la vie peut ainsi être décrite sous forme d'un « arbre évolutif », ou arbre
L'idée d'évolution peut déjà se trouver chez certains philosophes de l'antiquité (grecs, romains) ou
bien musulmans, mais ce n'est qu'à partir du XIXe siècle que de véritables théories proposant une explication
du phénomène de l'évolution des espèces ont été développées. Si la théorie du transformisme de Lamarck a
ouvert la voie, la révolution évolutionniste est arrivée avec Charles Darwin et son ouvrage De l'origine des
espèces (1859) dans lequel deux grandes idées, appuyés par des faits, émergent : l'unité et la diversité du
vivant s'explique par l'évolution, et le moteur de l'évolution adaptative est la sélection naturelle. En profonde
contradiction avec les idées philosophiques et religieuses dominantes de l'époque, De l'origine des espèces
obtient un important écho et convainc rapidement la majorité des biologistes de la réalité de l'évolution[1].
Avec la théorie synthétique de l'évolution qui regroupe notamment les idées de Darwin avec celles
de Mendel, l'évolution fait l'objet d'un large consensus scientifique sur ses fondements et ses mécanismes
depuis le milieu du XXe siècle. Dans la biologie contemporaine, si l'idée que les espèces évoluent ne fait plus
aucun doute, les détails des mécanismes qui permettent d'expliquer cette évolution font toujours l'objet de
recherches et sont parfois au cœur de controverses scientifiques et médiatiques, comme celle ayant opposé
Stephen Jay Gould et Richard Dawkins sur l'intérêt d'introduire la notion d'équilibres ponctués.
L'évolution est causée, d'une part par la présence de variations parmi les traits héréditaires d'une
population d'individus (mutations), et d'autre part par divers mécanismes qui vont modifier la fréquence de
certains traits héréditaires. Parmi ces mécanismes, la sélection naturelle désigne la différence de
propagation entre les traits héréditaires causée par leur effet sur la survie et la reproduction des individus : si
un certain trait héréditaire favorise les chances de survie et la reproduction, il s'ensuit mécaniquement que la
fréquence de ce trait augmente d'une génération à l'autre. Dans une population de taille finie, un trait peut
également être propagé ou éliminé par le fait de fluctuations aléatoires (dérive génétique). À l'échelle des
temps géologiques, l'évolution conduit à des changements morphologiques, anatomiques, physiologiques et
comportementaux des espèces.
Si l'on veut retranscrire ces concepts en systématique, il faut considérer la théorie cladistique, selon
laquelle les grades évolutifs (qui induisent une vision de l'évolution aujourd'hui obsolète [réf. nécessaire]) ne
sont plus pris en compte, en faveur des clades[2].
Du fait, entre autres, de ses implications sur l'origine de l'humanité, l'évolution a été, et reste
toujours, mal comprise et/ou, parfois, mal admise hors de la communauté scientifique. Dans les sociétés
occidentales, la théorie de l'évolution se heurte à une vive opposition de la part de certains milieux religieux
fondamentalistes, notamment pour son incompatibilité avec une interprétation littérale de la Bible. Ses
détracteurs se basent sur des analyses pseudo-scientifiques ou religieuses pour contredire l'idée même
d'évolution des espèces ou la théorie de la sélection naturelle.
• 1 Histoire de la théorie de l'évolution
• 2 Les arguments en faveur de l'évolution.
• 2.1 Stratégie de raisonnement
• 2.2 Indices morphologiques
• 2.3 Indices moléculaires
• 2.4 Indices comportementaux
• 2.5 Un exemple d'évolution à échelle de temps
humaine : Podarcis sicula
• 3 Les critiques de la théorie synthétique de l'évolution
• 4 Méthodes d'étude de l'évolution
• 4.1 La paléobiologie
• 4.2 L'analyse comparative des caractères
• 4.3 La génétique des populations
• 4.4 La modélisation
• 4.5 L'expérimentation
• 5 Mécanismes de l'évolution
• 5.1 L'évolution des populations
• 5.1.1 L'apparition de nouveaux caractères
• 5.1.2 Variabilité des individus au sein des
Histoire de la théorie de l'évolution [modifier]
Article détaillé : Histoire de la pensée évolutionniste.
Bien que les hommes cherchaient l'origine de la diversité du vivant dès la période antique, l'idée
d'évolution, c'est-à-dire de modification des espèces au cours du temps, a mis du temps à s'imposer. Si
l'idée d'une évolution de la vie est déjà présente chez quelques philosophes grecs[3], l'un de ceux qui a le
plus marqué le monde occidental[4], Aristote, avait une conception fixiste du vivant, et cette vision est restée
prédominante dans la pensée occidentale jusqu'au XVIIIe siècle. L'influence des religions monothéistes
abrahamiques est prédominante dans la diffusion de ces idées fixistes, sous une nouvelle forme : le
créationnisme. En effet les récits bibliques, en particulier ceux de la Genèse, prônent que toutes les espèces
vivantes ont été créées telles quelles et de manière synchrone par Dieu et qu'elles sont parfaites donc
immuables ; de plus, l'homme occuperait une place à part dans le vivant puisqu'il serait à l'image de Dieu et
serait moralement supérieur à toutes les autres espèces[3].
Durant le Moyen Âge, les avancées scientifiques en Europe occidentale deviennent limitées par la
dominance du fondamentalisme chrétien, qui prône une interprétation littérale des textes sacrés[5]. Bien que
les autorités religieuses condamnent fermement toute idée scientifique remettant en cause les écrits
bibliques, l'idée d'évolution se retrouve chez certains savants comme Jérôme Cardan[6] et Giulio Cesare
Vanini[7]. Parallèlement, l'idée d'évolution apparait dans le monde musulman, et l'on trouve dès le IXe siècle
non seulement l'idée que les espèces évoluent au cours du temps, mais aussi une première théorie
cherchant à expliquer cette évolution[8]. Au XIIIe siècle, le philosophe Nasir ad-Din at-Tusi propose,
plusieurs siècles avant Charles Darwin, la sélection des meilleurs et l'adaptation des espèces à leur
environnement[9]. Cependant, ces idées n'ont eu qu'une faible popularité, y compris au sein du monde
musulman[réf. souhaitée].
Au début du XVIIIe siècle, les idées fixistes alors prédominantes sont ébranlées par le
développement de la paléontologie et la découverte de fossiles de squelettes ne ressemblant à aucun
squelette actuel[10]. Pour concilier ces découvertes avec les textes bibliques, Georges Cuvier expose sa
théorie catastrophiste selon laquelle il y aurait eu une succession de créations divines entrecoupées
d'extinctions brutales au cours des temps géologiques[11]. Ils admet ainsi que les espèces terrestres n'ont
pas toujours été celles observées aujourd'hui, sans pour autant accepter l'évolution des espèces, et que les
6 000 ans estimés jusque là pour l'âge de la Terre sont trop courts pour y intégrer ces extinctions
Si l'idée d'évolution est réapparue au milieu du XVIIIe siècle avec Maupertuis et Buffon, la première
théorie véritablement scientifique considérant une évolution des espèces vivantes, le lamarckisme, est
fondée par le naturaliste Jean-Baptiste Lamarck.
Lamarck considère que les espèces peuvent se transformer selon deux principes :
1. La diversification, ou spécialisation, des êtres vivants en de multiples espèces, sous l'effet
des circonstances variées auxquelles ils sont confrontés dans des milieux variés et auxquelles ils
sont contraints de s'adapter en modifiant leur comportement ou leurs organes pour répondre à leurs
besoins, généralement désigné par « l'usage et le non-usage » ;
2. la complexification croissante de l'organisation des êtres vivants sous l'effet de la dynamique
interne propre à leur métabolisme.
Sa théorie a souvent été abusivement réduite à celle de la transmission des caractères acquis, qui
veut que les modifications acquises au cours de la vie d'un organisme soit héréditaires, mais en fait Lamarck
ne propose pas de théorie de l'hérédité de l'acquis (contrairement à ce que fera Darwin en 1868), il se
contente de reprendre les idées admises sur ce point depuis Aristote. Il n'en reste pas moins que Lamarck
est le premier à proposer une théorie expliquant les êtres vivant à partir de laquelle il tente de comprendre
l'évolution des espèces. La publication, en 1809, dans Philosophie zoologique, de sa théorie transformiste
entraine de virulents débats au sein de l'Académie des sciences car elle entre en contradiction avec les
idées en vigueur à l'époque et notamment le fixisme. Malgré de nombreuses critiques de la part des milieux
religieux et scientifique, et notamment de la part de Cuvier qui devient le principal opposant des
transformistes, les idées transformistes reçoivent une adhésion croissante à partir de 1825 et permettent de
rendre le débat naturaliste plus réceptif aux théories évolutionnistes[13] même si Lamarck ne verra jamais
ses travaux reconnus par la communauté scientifique[14].
Cependant, la première théorie satisfaisante permettant d'expliquer l'adaptation des espèces est
publiée en 1859 par Charles Darwin dans son livre De l'origine des espèces. Cette théorie, fondement de la
théorie actuelle de l'évolution, considère que, étant donné que tous les individus d'une espèce diffèrent au
moins légèrement, et que seule une partie de ces individus réussit à se reproduire, seuls les descendants
des individus les mieux adaptés à leur environnement participeront à la génération suivante. Ainsi, comme
les individus sélectionnés transmettent leurs caractères à leur descendance, les espèces évoluent et
s'adaptent en permanence à leur environnement. Il baptise du nom de sélection naturelle cette sélection des
individus les mieux adaptés[15]. Darwin n'avait cependant aucune idée du mécanisme permettant la
transmission des caractères[réf. souhaitée].
La découverte des lois de Mendel et de la génétique au début du XXe siècle bouleverse la
compréhension des mécanismes de l'évolution et donne naissance à la Théorie synthétique de l'évolution,
fondée entre autres par Theodosius Dobzhansky et Ernst Mayr. Cette théorie est une combinaison de la
théorie de la sélection naturelle proposée par Darwin et de la génétique mendellienne. Elle est à l'origine de
nouvelles méthodes dans l'étude de l'évolution, comme la génétique des populations ou la
modélisation[réf. nécessaire].
À partir de ce moment, la biologie de l'évolution intègre toutes les autres disciplines de la biologie, et
cherche aussi bien à retracer l'histoire évolutive du vivant qu'à théoriser et prouver les mécanismes en jeu
lors de l'évolution des espèces. La fin du XXe siècle est ainsi très prolifique sur le plan théorique. Plus
récemment, l'étude de l'évolution profite du développement de l'informatique et de la biologie moléculaire, et
notamment du séquençage qui permet le développement de la phylogénie par un apport très important de
données[réf. nécessaire].
La biologie de l'évolution est aujourd'hui une composante majeure de la biologie qui nourrit aussi
bien qu'elle se nourrit de toutes les autres disciplines[réf. nécessaire].

Les arguments en faveur de l'évolution. [modifier]

Stratégie de raisonnement [modifier]

Si on arrive à établir un lien de parenté entre deux espèces différentes, alors cela veut dire qu'une
espèce ancestrale s'est transformée en, au moins, une de ces deux espèces. Il y a donc bien eu évolution.
Un lien de parenté entre espèces fossiles ou actuels peut être mis en évidence par le partage d'au
moins un caractère homologue, c'est-à-dire provenant d'un ancêtre. Ces indices de parenté sont décelables
au niveau de la morphologie, au niveau moléculaire et parfois même, pour des espèces très proches, au
niveau du comportement.
Utilisation des fossiles
Archaefructus liaoningensis le plus ancien reste de fleur connu.
Il est en pratique impossible d'affirmer qu'une espèce fossile est l'ancêtre d'une espèce actuelle, car
il ne sera jamais garanti que l'espèce actuelle ne s'est pas différenciée à partir d'une autre espèce proche,
mais qui n'aurait pas été découverte. En effet, la conservation de restes d'espèces éteintes est un
événement relativement improbable surtout pour les périodes les plus anciennes. On peut seulement estimer
les liens de parenté, avec les autres espèces déjà connnues, actuelles ou fossiles. Par exemple le fossile de
fleur le plus ancien a été daté de 140 millions d'années. Cet organe est donc apparu sur Terre, il y a au
moins 140 millions d'années. Mais d'autres espèces proches, avec des fleurs, existaient aussi certainement
à cette époque. Personne n'est capable d'affirmer laquelle de ces espèces est l'ancêtre des plantes à fleur
actuelles. On ne cherchera que les relations de parenté, les relations d'ancêtre à descandant ne pouvant
jamais être reconstituées.
L'âge d'une espèce fossile, en revanche, indique l'âge minimum d'apparition des caractères qu'elle
possède. Il est alors possible de reconstruire l'histoire de l'évolution, en plaçant sur une échelle des temps
l'apparition des différents caractères. Les fossiles nous indiquent que l'ordre d'apparition des innovations
évolutives est tout à fait en accord avec l'idée d'une évolution, qui dans un schéma général, part de
structures simples vers des structures plus complexes. C'est aussi en accord avec une origine aquatique des
êtres vivants.

Indices morphologiques [modifier]

Squelette de baleine. En c figure le vestige de bassin[16].

Les « mains » des tétrapodes ont un même plan d'organisation, ce qui traduit une homologie.
• les baleines, animaux adaptés à la vie aquatique gardent une trace de leurs ancêtres
quadrupèdes par la présence d'os vestigials correspondant au bassin (ceinture pelvienne) ;
• Il y a des vestiges de pattes chez certains serpents (boas)[17];
• En observant l'aile d'un oiseau ou d'une chauve-souris, on retrouve aisément la structure
osseuse du membre antérieur de tout tétrapode ;
• les défenses à croissance continue des éléphants sont en fait homologues des incisives des
autres mammifères, dont l'homme ;
• les appendices masticateurs des arthropodes sont à l'origine des appendices locomoteurs
réduits (il en va de même apparemment pour les Onychophores) ;
• les membres des tétrapodes proviennent des nageoires de poissons ;
• dans le monde végétal, la présence d'une double membrane autour des plastes et la
présence d'un ADN circulaire à l'intérieur de ceux-ci trahissent une origine endosymbiotique

Indices moléculaires [modifier]

• Le support de l'information héréditaire est toujours l'ADN pour l'ensemble du vivant ;
• Le code génétique, code de correspondance entre l'ADN et les protéines est quasiment le
même chez tous les êtres vivants ;
• Le séquençage de l'ADN. fait apparaître de nombreuses régions étroitement proches donc
apparentés (gènes homologues: paralogues ou orthologue) qui codent des protéines aux fonctions
ou structures différentes mais assez proches (Exemple : les gènes qui codent les hémoglobines,

Indices comportementaux [modifier]

Chez certaines espèces de Lacertidés américains du genre Cnemidophorus, ou lézards à queue en
fouet, il n'existe plus que des femelles. Ces espèces pratiquent donc une reproduction asexuée. Cependant
des simulacres d'accouplements persistent : pour se reproduire une femelle monte sur une autre dans un
comportement similaire à celui des espèces sexuées. Ce comportement d'origine hormonale est à mettre en
relation avec une origine récente de ces espèces parthénogénétiques[18].
Un exemple d'évolution à échelle de temps humaine : Podarcis sicula [modifier]

Podarcis sicula. Des lézards des ruines déposés sur l'île de Prod Mrcaru en 1971 ont évolué en 36
ans de sorte à disposer d'un nouvel organe de digestion absent chez l'espèce d'origine : les valves cæcales.
Introduit en 1971 par l'équipe du professeur Eviatar Nevo sur l'île dalmate de Prod Mrcaru en mer
Adriatique, le lézard Podarcis sicula connu en France sous le nom de « lézard des ruines », y a été
abandonné à lui-même durant près de quatre décennies, l'accès à l'île ayant été interdit par les autorités
yougoslaves, puis par les conflits liés à l'éclatement de ce pays. En 2004, une équipe scientifique dirigée par
Duncan Irschick et Anthony Herrel put revenir sur l'île et découvrit que Podarcis sicula avait évolué en 36
ans, soit environ trente générations, de façon très significative. Le lézard a grandi, sa mâchoire est devenue
plus puissante, et surtout il a changé de régime alimentaire : d'insectivore il est devenu herbivore, et des
valves cæcales sont apparues au niveau des intestins, ce qui lui permet de digérer les herbes... Cette
découverte confirme, s'il en était encore besoin, que l'évolution n'est pas une théorie parmi d'autres, mais un
phénomène biologique concrètement observable, et pas seulement chez les virus, les bactéries ou les
espèces domestiquées[19].

Les critiques de la théorie synthétique de l'évolution [modifier]

Article détaillé : Historique des critiques des théories de l'évolution.
Ceux-ci peuvent être rapidement classés en diverses origines :
• Critiques idéologiques.
• Critiques scientifiques (notamment par rapport aux fossiles).
• Critiques religieuses (créationnisme et intelligent design).

Méthodes d'étude de l'évolution [modifier]

La paléobiologie [modifier]
Articles détaillés : Paléontologie et Paléogénétique.
La paléobiologie, étude de la vie des temps passés, permet de reconstituer l'histoire des êtres
vivants. Cette histoire donne aussi des indices sur les mécanismes évolutifs en jeu dans l'évolution des
espèces. La paléontologie s'occupe plus particulièrement des restes fossiles des êtres vivants. La
paléogénétique, science récente, s'intéresse au matériel génétique ayant survécu jusqu'à aujourd'hui[20].
Ces deux approches sont limitées par la dégradation du matériel biologique au cours du temps. Ainsi, les
informations issues des restes sont d'autant plus rares que l'être vivant concerné est ancien. De plus,
certaines conditions sont plus propices que d'autres à la conservation du matériel biologique. Ainsi, les
environnements anoxiques ou très froids entravent la dégradation des restes. Les restes vivants sont donc
lacunaires et sont bien souvent insuffisants pour retracer l'histoire évolutive du vivant.

L'analyse comparative des caractères [modifier]

Articles détaillés : Génétique évolutive du développement et Phylogénie.
Tous les êtres vivants actuels étant issus d'un même ancêtre commun, ils partagent des
caractéristiques héritées de cet ancêtre. L'analyse des ressemblances entre êtres vivants donne de
nombreuses informations sur leurs relations de parenté, et permet de retracer l'histoire évolutive des
espèces. La phylogénie est la discipline scientifique qui cherche à retracer les relations entre êtres vivants
actuels et fossiles à partir de l'analyse comparative des caractères morphologiques, physiologiques ou
moléculaires. L'analyse comparative permet de retracer l'histoire évolutive des différents caractères dans les
lignées du vivant. L'évolution des caractères ne suit pas nécessairement celle des espèces, certains
caractères (dits convergents) peuvent être apparus plusieurs fois de manière indépendante dans différentes
L'évolution des caractères et des lignées peut être associée à des évènements géologiques ou
biologiques marquant l'histoire de la Terre, ce qui permet de proposer des hypothèses sur les mécanismes à
l'origine de l'évolution des espèces.
La nature des caractères pouvant être analysés est extrêmement diverse, et il peut s'agir aussi bien
de caractères morphologiques (taille, forme ou volume de différentes structures), anatomiques (structure,
organisation des organes), tissulaires, cellulaires ou moléculaires (séquences protéiques ou nucléiques).
Ces différents caractères apportent des informations diverses et souvent complémentaires. Actuellement, les
caractères moléculaires (en particulier les séquences d'ADN) sont privilégiées, du fait de leur universalité, de
leur fiabilité et du faible coût des technologies associées. Ils ne peuvent cependant pas être utilisés lors de
l'étude de fossiles pour lesquels seuls les caractères morphologiques sont en général informatifs.

La génétique des populations [modifier]

Article détaillé : Génétique des populations.

La modélisation [modifier]
La modélisation en biologie de l'évolution se base sur les mécanismes de l'évolution mis en évidence
pour mettre en place des modèles théoriques. Ces modèles peuvent produire des résultats qui dépendent
des hypothèses de départ de ce modèle, ces résultats pouvant être comparés à des données réellement
observées. On peut ainsi tester la capacité du modèle à refléter la réalité, et, dans une certaine mesure, la
validité de la théorie sous-jacente à ce modèle.
Les modèles dépendent souvent de paramètres, lesquels ne peuvent pas toujours être déterminés a
priori. La modélisation permet de comparer les résultats du modèles et ceux de la réalité pour de
nombreuses valeurs différentes de ces paramètres, et ainsi déterminer quelles sont les combinaisons de
paramètres qui permettent au modèle décrire au mieux la réalité. Ces paramètres correspondent souvent à
des paramètres biologiques, et on peut ainsi estimer à partir du modèle certains paramètres biologiques
difficile à mesurer. La justesse de l'estimation de ces paramètres dépend cependant de la validité du modèle,
laquelle est parfois difficile à tester.
La modélisation permet enfin de prédire certaines évolutions à venir, en utilisant les données
actuelles comme données de départ du modèle.

L'expérimentation [modifier]
Article détaillé : Évolution expérimentale.
L'évolution expérimentale est la branche de la biologie qui étudie l'évolution par de réelles
expériences, à l'inverse de l'étude comparative des caractères, qui ne fait que regarder l'état actuel des êtres
vivants. Les expériences consistent généralement en l'isolement d'une ou plusieurs espèces dans un milieu
biologique contrôlé. On laisse alors ces espèces évoluer pendant un certain temps, en appliquant
éventuellement des changements contrôlés de conditions environnementales. On compare enfin certaines
caractéristiques des espèces avant et après la période d'évolution.
L'évolution expérimentale permet non seulement d'observer l'évolution en cours, mais aussi de
vérifier certaines prédictions énoncées dans le cadre de la théorie de l'évolution, et tester l'importance
relative de différents mécanismes évolutifs.
L'évolution expérimentale ne peut étudier que des caractères évoluant rapidement, et se limite donc
à des organismes se reproduisant rapidement, notamment des virus ou des unicellulaires, mais aussi
certains organismes à génération plus longue comme la drosophile ou certains rongeurs.
Un exemple : l'expérience de Luria et Delbrück.

Mécanismes de l'évolution [modifier]

Article détaillé : Théorie synthétique de l'évolution.
L'évolution des populations [modifier]

L'évolution des caractères dans les populations: diversité, sélection et transmission

Parce que les individus d'une population possèdent des caractères héritables différents, et que seule
une partie de ces individus accède à la reproduction, les caractères les plus adaptés à l'environnement sont
préférentiellement conservés par la sélection naturelle. De plus, le hasard de la reproduction sexuée rend
partiellement aléatoire les caractères qui seront transmis, par effet de dérive génétique. Ainsi, la proportion
des différents caractères d'une population varie d'une génération à l'autre, conduisant à l'évolution des

L'apparition de nouveaux caractères [modifier]

Cela se produit par mutation et recombinaison génétique, ou remaniement chromosomique. Mais
cela ne se déroule que dans un individu, pas dans l'espèce entière. Il faut, pour que ce nouveau caractère se
répande, l'effet de la sélection naturelle et/ou de la dérive génétique.

Variabilité des individus au sein des populations [modifier]

Articles détaillés : Diversité génétique, Mutation et Reproduction sexuée.
Tous les individus d'une espèces sont uniques et diffèrent les uns des autres. Ces différences sont
observables à toutes les échelles, du point de vue morphologique jusqu'à l'échelle moléculaire. Cette
diversité des populations a deux origines principales: les individus sont dissemblables parce qu'ils ne
possèdent pas la même information génétique et parce qu'ils ont subi des influences environnementales
La diversité génétique se manifeste par des variations locales de la séquence d'ADN, formant
différents variants de la même séquence appelés allèles. Cette variabilité a plusieurs origines. Des allèles
peuvent être formés spontanément par mutation de la séquence d'ADN. Par ailleurs, la reproduction sexuée
contribue à la diversité génétique des populations de deux manières: d'une part, la recombinaison génétique
permet de diversifier les combinaisons d'allèles réunies sur un même chromosome. D'autre part, une partie
du génome de chaque parent est sélectionnée aléatoirement pour former un nouvel individu, dont le génome
est par conséquent unique.
La diversité issue de l'environnement s'acquiert tout au long de l'histoire de l'individu, depuis la
formation des gamètes jusqu'à sa mort. L'environnement étant unique à chaque endroit et à chaque moment,
il exerce des effets unique sur chaque individu, et ce à toutes les échelles, de la morphologie jusqu'à la
biologie moléculaire. Ainsi, deux individus possédant la même information génétique (c'est par exemple le
cas pour les vrais jumeaux) sont tout de même différents. Ils peuvent notamment avoir une organisation et
une expression différente de l'information génétique.

L'hérédité [modifier]
Articles détaillés : Hérédité, ADN, Reproduction, Réplication de l'ADN et Épigénétique.
Les êtres vivants sont capables de se reproduire, transmettant ainsi une partie de leurs caractères à
leurs descendants. On distingue la reproduction asexuée, ne faisant intervenir qu'un individu, de la
reproduction sexuée pendant laquelle deux individus mettent en commun une partie de leur matériel
génétique, formant ainsi un individu génétiquement unique.
mitochondries, n'est transmis que par une partie des individus de l'espèce (les femelles chez les

La transmission des caractères acquis, une hypothèse non totalement rejetée [modifier]
Article détaillé : Transmission des caractères acquis.
La théorie synthétique de l'évolution, paradigme dominant actuel, se fonde sur un déterminisme
génétique intégral et écarte donc toute transmission héréditaire de caractères acquis au cours de la vie de
l'individu. Néanmoins de plus en plus de travaux scientifiques remettent en cause ce modèle et rétablissent
pour partie l'idée d'une transmission héréditaire de caractères acquis que défendait le lamarckisme[21].
Tout d'abord, certains caractères dits épigénétiques concernent la structure et l'organisation des
génomes sont transmis par les parents en même temps que les molécules d'acide nucléique elles-mêmes.
De plus, la mère fournit l'environnement cytoplasmique de la cellule-oeuf du descendant, et transmet ainsi un
certain nombre de caractéristiques cellulaires à l'enfant. Des modifications épigénétiques conservées dans la
lignée germinale sont désormais décrites chez plusieurs espèces. Chez les plantes il existe une corrélation
entre le niveau d'expression d'un gène et sa méthylation. Pareillement, chez les mammifères nous
témoignons de la méthylation d'une séquence transposable qui est insérée à proximité d'un gène particulier.
Le degré de méthylation d'un transposon pouvant enfin moduler l'expression du gène dans lequel il s'est
inséré[22]. L'étude de l'épigénétique, longtemps délaissée, connait un grand essor depuis la fin du
séquençage de nombreux génomes, dont celui de l'homme.
Ainsi, Une étude de 2009 du MIT affirme mettre en évidence une hérédité de certains caractères
acquis chez des rongeurs[23]. Par ailleurs, l’obésité serait non pas uniquement un effet direct touchant les
individus atteints eux-mêmes mais également un effet transgénérationnel. Des données chez l'homme et
chez l'animal semblent montrer que les effets d'une sous-alimentation subies par des individus pourraient en
effet être transmises aux descendants.

La dérive génétique [modifier]

Article détaillé : dérive génétique.
Simulation informatique de l'évolution de la fréquence d'un allèle neutre au cours du temps dans une
population de 10 (en haut) ou 100 individus (en bas). Chaque courbe représente une simulation différente,
les différences illustrant l'effet du hasard (dérive génétique). Les fluctuations de la fréquence de l'allèle sont
plus importantes lorsque la population est de taille réduite, et la fixation (fréquence égale à 1) ou la perte
(fréquence égale à 0) d'un allèle est alors plus rapide.
Lors de la reproduction sexuée, la transmission des caractères (notamment des allèles) comporte
une grande part de hasard due à la recombinaison homologue, et au brassage génétique. Ainsi, on observe
une variation aléatoire des fréquences alléliques d'une génération à l'autre, appelée dérive génétique. La
dérive génétique génère donc une composante aléatoire dans l'évolution des populations. Ainsi, deux
populations d'une même espèce n'échangeant pas de matériel génétique vont diverger jusqu'à former, si le
temps d'isolement génétique est suffisant, deux espèces différentes. La dérive génétique est donc un des
moteurs de la spéciation.
L'effet de la dérive génétique est particulièrement visible lorsqu'un faible nombre d'individus est à
l'origine d'une population beaucoup plus nombreuse. C'est le cas lorsque se forme un goulot d'étranglement
c'est-à-dire qu'une population est décimée et se reconstitue, ou lorsque quelques individus d'une population
migrent pour aller coloniser un nouvel espace et former une nouvelle population (effet fondateur). Lorsqu'un
tel évènement se produit, un allèle même faiblement représenté dans la population de départ peut se
retrouver en forte proportion dans la population nouvellement formée sous le simple effet d'un hasard dans le
tirage des individus à l'origine de la nouvelle population. Inversement, un allèle fortement représenté peut ne
pas être tiré, et disparaît de la nouvelle population. Par ailleurs, la formation d'une nouvelle population à
partir d'un faible nombre d'individu a pour effet d'augmenter la consanguinité dans la population et augmente
le pourcentage d'homozygotie, ce qui fragilise la population.
La sélection naturelle [modifier]
Article détaillé : Sélection naturelle.
Dans la très grande majorité des espèces, le nombre de cellules-œuf produits est bien plus grand
que le nombre d'individus arrivant à l'âge de la maturité sexuelle et parmi ceux-ci, une partie seulement
accède à la reproduction. Ainsi, seule une partie des individus formés se reproduit à la génération suivante. Il
existe donc une sélection des individus perpétuant l'espèce, seuls les individus n'étant pas éliminé par les
conditions environnementales pouvant se reproduire. Cette sélection a été baptisée sélection naturelle.
Comme il existe une variabilité au sein des espèces, les individus possédant des caractères
différents, et qu'une partie de ces caractères sont héréditaires, les caractères permettant à l'individu de
survivre et de mieux se reproduire seront préférentiellement transmis à la descendance, par rapport aux
autres caractères. Ainsi la proportion des caractères au sein des espèces évolue au cours du temps.
La sélection naturelle peut prendre des formes très variées. La sélection utilitaire est une élimination
des individus les moins capables de survivre et les moins féconds, alors que la sélection sexuelle conserve
préférentiellement les individus les plus aptes à rencontrer un partenaire sexuel. Bien que ces sélections
soient complémentaires, on observe souvent des conflits, chaque forme de sélection pouvant favoriser
l'évolution d'un caractère dans un sens différent.
Il est parfois observé une sélection d'individus qui favorisent la survie ou la reproduction d'individus
qui leurs sont ou non apparentés, comme c'est le cas chez les insectes eusociaux ou lorsqu'un individu se
sacrifie pour permettre la survie de son groupe ou de sa descendance. En sociobiologie, ces comportements
altruistes s'expliquent notamment par les théories controversées de la sélection de parentèle, de la sélection
de groupe et de l'altruisme réciproque. La sélection de parentèle prédit qu'il peut être plus avantageux pour
un individu de favoriser beaucoup la reproduction d'un individu apparenté (donc avec lequel il partage des
caractères) que de se reproduire un peu ou pas du tout, la sélection de groupe repose sur le même principe
mais du point de vue du groupe et pourrait expliquer certains actes chez l'homme comme les guerres ou la
xénophobie, l'altruisme réciproque se penche sur les cas d'altruisme entre individus non-apparentés et induit
une contribution réciproque dont l'aide donnée en retour peut être différé dans le temps.
Enfin, la sélection artificielle n'est qu'une forme de sélection naturelle exercée par l'homme.

Conséquences évolutives [modifier]

Adaptation des espèces [modifier]

Articles détaillés : Adaptation et Neutralisme.
En conséquence de la sélection naturelle, les espèces conservent préférentiellement les caractères
les plus adaptés à leur environnement, et sont donc de mieux en mieux adaptées à leur environnement. Les
pressions de sélection en jeux dans cette adaptation sont nombreuses et concernent tous les aspects de
l'environnement, des contraintes physiques jusqu'aux espèces biologiques interagissantes.
L'adaptation de plusieurs espèces différentes sous l'effet des mêmes pressions environnementales
peut conduire à l'apparition répétée et indépendante du même caractère adaptatif chez ces espèces, par un
phénomène de convergence évolutive. Par exemple, chez les mammifères les cétacés et les siréniens ont
tout deux développé des nageoires, de manière indépendante. L'évolution de ces nageoires montre une
adaptation convergente à la vie aquatique.
Cependant, l'effet de la sélection naturelle est réduit par celui de la dérive génétique. Ainsi, un
caractère avantageux pourra ne pas être sélectionné à cause de l'inertie donnée par la dérive.

Apparition et disparition des espèces [modifier]

Articles détaillés : Spéciation et Extinction des espèces.
L'évolution d'une population sous l'effet du hasard et des contraintes environnementales peut aboutir
à la disparition de la population et éventuellement de l'espèce à laquelle elle appartient. Inversement, deux
populations peuvent s'individualiser au sein d'une même espèce jusqu'à former deux espèces distinctes par
un processus nommé spéciation.

Controverses sur les mécanismes de l'évolution [modifier]

Cet article ne cite pas suffisamment ses sources (décembre 2009).
Si vous connaissez le thème traité, merci d'indiquer les passages à sourcer avec
{{Référence souhaitée}} ou, mieux, incluez les références utiles en les liant aux notes de
bas de page. (Modifier l'article)
L'évolution et ses mécanismes sont encore largement étudiés aujourd'hui, et de nombreux points sur
les mécanismes de l'évolution ne sont pas éclaircis. Certaines questions déjà soulevées par Charles Darwin
n'ont d'ailleurs toujours pas de réponse certaine.
Une des grandes questions de la théorie de l'évolution est l'origine des rangs taxinomiques
supérieurs à celui de l'espèce. En outre, la manière dont sont apparus les 33 embranchements animaux,
issus de l'explosion cambrienne, pose encore problème. Ainsi, la théorie gradualiste estime que les
changements interviennent de manière progressive au cours de l'évolution, alors que la théorie des
équilibres ponctués défend qu'il existe des sauts évolutifs majeurs.
La transmission des caractères acquis, complètement délaissée depuis la découverte des lois de
l'hérédité, est réactualisée par la découverte des phénomènes épigénétiques. Dès lors, l'importance de cette
transmission de caractères non hérités des parents dans l'évolution des espèces doit se poser. Cependant,
notre connaissance des mécanismes épigénétiques est encore trop faible pour pouvoir répondre à cette
question. En outre, peu d'études sur le rôle de l'épigénétique dans l'évolution ont été réalisées à l'heure
Il a été longtemps admis que l'évolution s'accompagnait d'un accroissement de la complexité des
êtres vivants. Cependant, cette idée, largement influencée par l'anthropocentrisme, est fortement débattue
aujourd'hui. La complexité n'ayant pas de définition précise à l'heure actuelle, il est difficile de vérifier une
éventuelle augmentation de complexité. Par ailleurs, lorsque cette idée est admise, les origines de cette
augmentation de complexité sont, elles aussi, source de controverse. En fait, tout cela à déjà été clairement
expliqué par Lamarck.

Histoire évolutive du vivant [modifier]

L'origine de la Vie se situerait vers - 3.8 milliards d'années. Il s'agissait probablement d'organismes
procaryotes unicellulaires. On suppose un ancêtre unique à tous les êtres vivants (LUCA). À partir de cet
ancêtre se sont diversifiées les différentes formes de Vie.
Articles détaillés : Histoire évolutive du vivant et Arbre phylogénétique du vivant.
Évolution et sociétés humaines [modifier]

Évolution et agriculture [modifier]

Article détaillé : Sélection artificielle.
L'homme a su très vite utiliser la variabilité des populations à son profit : l'évolution dirigée par
l'homme, ou sélection artificielle, à cause de la sélection par les éleveurs et les cultivateurs, se produit
depuis des millénaires. Il avait été remarqué depuis longtemps que les animaux d'élevage héritaient, dans
une certaine mesure, de caractéristiques de leurs parents et nul n'aurait songé à utiliser ses bêtes les plus
malingres pour la reproduction. D'ailleurs, Darwin utilise de nombreuses observations issues de la sélection
des plantes et des animaux en agriculture pour étayer ses idées. Ainsi, l'homme peut créer une sélection dite
artificielle sur son environnement, volontairement pour des raisons économiques, ou involontairement via la
pression de chasse, cueillette ou pêche)[24].

Évolution et informatique [modifier]

Article détaillé : Algorithme évolutionniste.
L'efficacité du processus de sélection naturelle a inspiré la création d'algorithmes évolutionnistes
(comme les algorithmes génétiques) en informatique. Ces algorithmes heuristiques modélisent plusieurs
caractéristiques de l'évolution biologique (en particulier les mutations et les recombinaisons) pour trouver
une solution satisfaisante à un problème trop complexe pour être abordé par d'autres méthodes.
Eugénisme [modifier]
Articles détaillés : Darwinisme social et Eugénisme.
Article détaillé : Évolutionnisme (anthropologie).
La pensée évolutionniste s'est notamment propagée au sein de l'anthropologie évolutionniste au
XIXe siècle. Pour les anthropologues de cette époque, l'espèce humaine ne fait qu'une, et donc, chaque
société suit la même évolution, qui commence à l'état de « primitif » pour arriver jusqu'au modèle de la
civilisation occidentale. Cette théorie a été très fortement remise en question. En effet, elle ne correspond
pas à la réalité historique observée (les civilisations suivent des « chemins » divergents, ne poursuivent pas
les mêmes « objectifs », et la civilisation occidentale, qui devrait pourtant constituer le stade ultime de
l'évolution, continue pourtant à vivre de profondes mutations.) et est douteuse d'un point de vue éthique
(considérant notre société occidentale comme l'aboutissement ultime de la civilisation). À l'inverse de ce qui
était pratiqué jusqu'au milieu du XXe siècle, les approches modernes de l'anthropologie évolutionniste
privilégient une méthodologie précise (confrontant des sources multiples, s'inspirant des outils d'analyse
quantitative des sciences sociales, tentant de se départir de l'ethnocentrisme) et s'appuie sur des théories
plus élaborées que l'évolutionnisme simpliste des débuts. Théories inspirées non seulement par la biologie
de l'évolution moderne mais aussi par la modélisation mathématique et informatique et parfois enrichies par
les connaissances contemporaines en psychologie.

La psychologie évolutionniste [modifier]

Article détaillé : Psychologie évolutionniste.
évolutionniste. Même si Darwin avait déjà émis l'idée que la sélection naturelle a pu façonner aussi bien des
caractères anatomiques que psychologiques, cette discipline s'est véritablement formalisée au début des
années 1990 dans le cadre conceptuel des sciences cognitives. Depuis, la psychologie évolutionniste est au
centre d'une intense controverse scientifique qui tient à de multiples raisons : difficulté méthodologique à
établir une histoire évolutive des comportements qui ne sont pas des objets matériels, résistance
intellectuelle à envisager l'esprit humain comme en partie déterminé par l'évolution, utilisation simpliste et
abusive des théories évolutionnistes, médiatisation et déformation auprès du grand public des
problématiques scientifiques... Dans le milieu scientifique toutefois, la psychologie évolutionniste fait
désormais partie des paradigmes scientifiques valides.

Evolutionnisme et religions [modifier]

La théorie évolutionniste est-elle compatible avec la croyance en Dieu ? En fait, Ernst Mayr dit à ce
sujet : « Il me semble évident que Darwin a perdu la foi un an sinon deux, avant de formuler sa théorie de la
sélection naturelle (sur laquelle il a sans doute travaillé plus de dix ans). Par conséquent, il n'est pas fondé
d'avancer que la biologie et l'adhésion à la théorie de la sélection naturelle risquent de vous éloigner de
Le biologiste Richard Dawkins, dans son ouvrage Pour en finir avec Dieu (2008), pense que la
sélection naturelle est « supérieure » à l'« hypothèse de Dieu » qu'il qualifie d'« improbabilité statistique », et
défend l'athéisme.
Le biologiste Kenneth R. Miller (en) estime que la pensée évolutionniste n'est pas forcément
incompatible avec la foi en un Dieu[26]. Pour lui les écrits de la Bible sont des métaphores.
L'évolution est encore aujourd'hui rejetée par certains milieux religieux, tenants du créationnisme,
surtout protestants.
La position de l'Église Catholique sur ce sujet est plus nuancée, tout en maintenant l'innerance de la
Bible[27], une microévolution au sein des espèces semble aujourd'hui « plus qu'une hypothèse ». Elle
déclare que Dieu est le seul créateur, qu'Il a créé le monde par amour, mais que l'esprit ne peut pas être le
fruit d'une évolution de la matière[28].

Aspects politiques et judiciaires [modifier]

Les polémiques ont débordé, depuis les années 1990, le simple cadre du débat public, notamment
aux États-Unis.
Dans certains États, les tenants du créationnisme ont essayé de rendre obligatoire son
enseignement dans les écoles publiques, en tant que « théorie scientifique concurrente » de celle de
l'évolution. Cependant ces mesures ont été déclarés anticonstitutionnelles vis-à-vis du premier amendement
sur la liberté d'expression, du fait du caractère religieux de cette théorie. Devant ces tentatives, des
scientifiques ont ironiquement demandé à ce que soit aussi enseigné le pastafarisme (qui a été inventé à
cette occasion).
Un nouveau concept est apparu dans la mouvance créationniste, baptisé dessein intelligent
(« Intelligent Design »), qui affirme que « certaines caractéristiques de l'Univers et du monde vivant sont
mieux expliquées par une cause intelligente, plutôt que par des processus aléatoires tels que la sélection
naturelle[29] ». Cette thèse est présentée comme une théorie appuyée par des travaux scientifiques, et ne
nie pas l'existence de tout phénomène évolutif. La justice américaine, s'appuyant sur les travaux
scientifiques, a cependant jugé (voir Kitzmiller v. Dover Area School) que cette thèse était de nature
religieuse et non scientifique, et que les promoteurs de l’ Intelligent Design n'explicitaient pas cette « cause
intelligente » afin de contourner le problème juridique et d'échapper au qualificatif religieux. D'autres groupes
utilisent les arguments de l’Intelligent Design, avec diverses attributions pour la « cause intelligente », par
exemple des extraterrestres.

Notes et références [modifier]

1. ↑ Darwin n'utilise pas le mot évolution dans son œuvre, puisque ce terme n'est introduit que
dans les années 1870. Cf. Gould (1997) : 33-37, Laurent (2001) : 17.
2. ↑ W. Hennig, Phylogenetic Systematics, Illinois University Press, 1966, traduit par D. Dwight
Davis & R. Zangerl.
3. ↑ a et b Barbara Cassin & al., L'animal dans l'Antiquité [lire en ligne [archive]], Centre National
de Recherche Scientifique, éd. Vrin, 1997, 618 p., (ISBN 2711613232).
4. ↑ Jean-Philippe Omotunde, Platon et Aristote : les deux piliers de la pensée
occidentale [archive], Institut Africamaat, (page consultée le 4 juillet 2008).
5. ↑ Jean Chaline, Quoi de neuf depuis Darwin ?, éd. Ellipses, 2006 (ISBN 978-2-7298-3100-
6. ↑ Jérôme Cardan définit une théorie de l'évolution dans son ouvrage De Subtilitate Rerum
en 1551.
7. ↑ Giulio Cesare Vanini est brûlé vif en 1619 pour avoir notamment déclaré que l'homme et le
singe pouvaient être des parents.
8. ↑ Mehmet Bayrakdar (The Islamic Quarterly Third Quarter, 1983). "Al-Jahiz And the Rise of
Biological Evolutionism [archive]", Londres.
9. ↑ Farid Alakbarli, « A 13th-Century Darwin? », dans Azerbaijan International, vol. 9.2, 2001,
p. 48-49
10.↑ Laurent Dubois, [pdf] Histoire de la paléontologie, Darwin et Théorie de
l'Evolution [archive], Gé, (page consultée le 4 juillet 2008).
11.↑ Lapierre, S., « Éléments de théorie de l'évolution [archive] » sur [archive]. Consulté le 19 octobre 2008
12.↑ Serge Lapierre, Éléments de théorie de l'évolution [archive], Collège de Bois de Boulogne
- Département de philosophie, (page consultée le 25 octobre 2008).
13.↑ Hélène Blais, « Lamarck, genèse et enjeux du transformisme, 1770-1830 », La Revue
pour l’histoire du CNRS [lire en ligne [archive]], n°7 - Novembre 2002, mis en ligne le 6 mars 2006.
Consulté le 7 juillet 2008.
14.↑ Benoît Virole, Le voyage intérieur de Charles Darwin: essai sur la genèse psychologique
d'une œuvre scientifique [lire en ligne [archive]], p. 124-126, éd. des archives contemporaines, 2000,
144 p. (ISBN 9057090171)
15.↑ Darwin, C., De l'Origine des espèces, Flammarion, 1859, 1997
16.↑ L. Bejder, B.K. Hall, « Limbs in whales and limblessness in other vertebrates: mechanisms
of evolutionary and developmental transformation and loss », dans Evol. Dev., vol. 6, no 4, Nov.-Dev.
2002, p. 445-58.
17.↑ (en)Vestigials Organs, A Snake—With Legs ! American Museum of Natural
History [archive] Site du muséum d'histoire naturelle américain.
18.↑ Serotonergic modulation of male-like pseudocopulatory behavior in the parthenogenetic
whiptail lizard, Cnemidophorus uniparens Brian George Dias et David Crews Hormones and
Behavior Volume 50, Issue 3, Septembre 2006, 401-409.
19.↑ Voir site : [pdf](en)PNAS, vol. 105, n°12, pages 4792-4795 [archive] (25 mars 2008).
20.↑ Eva-Maria Geigt, « L'émergence de la paléogénétique », dans Biofutur, no 164, Février
1997, p. 28-34 (ISSN 0294-3506)
21.↑ La théorie de l'évolution en évolution ? Hominidé [archive]
22.↑ Claudine Junien, « Obésité et diabète de type 2 : L'hypothèse de la transmission
épigénétique », dans Cahiers de nutrition et de diététique, vol. 37, no 4, 2002, p. 261-272
(ISSN 0007-9960) [ texte intégral [archive] (page consultée le 11/03/2009) ]
23.↑ [archive]
24.↑ Chris T. Darimonta et al. ; Human predators outpace other agents of trait change in the
wild ; Ed : Gretchen C. Daily, Stanford University, Stanford, CA, PNAS, approuvé le 21 nov 2008
(reçu pour relecture le 15 septembre 2008) (Lire l'article [archive])
25.↑ Biologie Campbell, De Boeck Université ISBN 2-8041-2084-8 p.417
26.↑ Kenneth Miller À la recherche du Dieu de Darwin, édition Sciences et quête de sens, 2000
16_fonti-pentateuco_fr.html [archive]
28.↑ [archive]
29.↑ « The theory of intelligent design [...] holds that certain features of the universe and of
living things are best explained by an intelligent cause rather than an undirected process such as
natural selection » — Intelligent Design Network, Inc. [archive]
Voir aussi [modifier]
Ouvrages sur le sujet [modifier]
Lorsqu'il y a deux dates, la première est • Jacob, F. (1981) Le Jeu des
celle de la première parution, dans la langue possibles, Fayard.
d'origine. • Kropotkine, P (2001, éd. or.
• Brondex, F. (1999) Évolution : 1902) L'entraide : un facteur de
synthèse des faits et théories, Dunod. l'évolution, Ecosociété.
• Buican, D. (1989) La Révolution de • Laurent, G. (2001). La
l'évolution, PUF. Naissance du transformisme. Lamarck
entre Linné et Darwin, Vuibert (Paris) et
• Buican, D. (1997) L'Évolution et les
ADAPT (Paris) : 151 p. (ISBN 2-7117-
théories évolutionnistes, Masson.
• Buican, D. (2008) L'odyssée de
• Dominique Lecourt, (1992, 3e
l'évolution, Ellipses
éd. « Quadrige » 1998), L’Amérique
• Chapouthier, G. (2001) L'homme, entre la Bible et Darwin, suivi de
ce singe en mosaïque, Odile Jacob Intelligent design : science, morale et
• Combes, C. (2006) Darwin, politique, PUF.
dessine-moi les hommes • Lecourt, D. dir. (1999, 4e réed.
• Darwin, C. (1997, éd. or. 1859) « Quadrige » 2006), Dictionnaire
L'Origine des espèces, Flammarion. d’histoire et philosophie des sciences,
• Charles Darwin. Origines - Lettres PUF.
choisies 1828-1859 (2009), introduction et • Le Guyader, H., dir. (1998)
édition française dirigée par Dominique L'évolution, Belin/Pour la Science.
Lecourt, préface S. J. Gould, éditions • Le Guyader, H. et Lecointre, G.,
Bayard, (ISBN 978-2-227-47843-5). Classification phylogénétique du vivant,
• David, P. & Samadi, S. (2000) La Belin (Paris) : 560 p. (ISBN 2-7011-
Théorie de l'évolution, Flammarion. 4273-3)
• Dawkins, R. (1982) The Extended • Lehman, J.-P. (1973) Les
Phenotype, Oxford University Press. preuves paléontologiques de l'évolution,
• Dawkins, R. (1986, 1989) PUF.
L'Horloger aveugle, Éditions Robert Laffont. • Lodé, T. (2006) La guerre des
• Dawkins, R. (1996) Climbing Mount sexes chez les animaux Odile Jacob,
Improbable, Norton (anglophone). Paris. (ISBN 2-7381-1901-8)
• Dawkins, R. (1976, 1996) Le Gène • Marchand, D. (2002) Les
égoïste, Odile Jacob. merveilles de l'évolution, P.U. Dijon.
• Dennett, D. (2000) Darwin est-il • Mayr, E. (1989) The Growth of
dangereux ?, Odile Jacob. Biological Thought: Diversity, Evolution
• Devillers, C. & Tintant, H. (1996) and Inheritance, Ed. Cambridge, Harvard
Questions sur la théorie de l'évolution, University Press — traduit en français
PUF. sous le titre de Histoire de la biologie.
• Dorléans, P. (2003) Il était une fois Diversité, évolution et hérédité, Fayard
l'évolution, Ellipses. (1989) : 894 p. (ISBN 2213018944).
• Futuyma, D.J (1997) Evolutionary • Mayr, E. (2004) What makes
Biology, Sinauer Associates. biology unique? Considerations on the
• Gould, S. J. (1982) Le Pouce du Autonomy of a Scientific Discipline, Ed.
New York, Cambridge University Press —
panda, Grasset. traduit en français sous le titre de Ernst
• Gould, S. J. (1991) La Vie est belle, Après Darwin. La biologie, une science
Le Seuil. pas comme les autres, Dunod (2006) :
• Gould, S. J. (1997) L'Éventail du 237 p. (ISBN 2-10-049560-7).
vivant, Le Seuil. • Pichot, A. (1993) Histoire de la
• Gould, S. J. (1997). Darwin et les notion de vie, éd. Gallimard, coll. TEL.
grandes énigmes de la vie. Réflexions sur • Ridley (1998) Evolution
l'histoire naturelle. 1, S 43, Seuil (Paris), Biologique, Ed. De Boeck (traduction
collection Point Science : 311 p. française).
• Gould, S. J. (2000) Et Dieu dit : • Tort, P. (1996) Dictionnaire du
Que Darwin soit ! : Science et religion, darwinisme et de l’évolution, Ed. Paris,
enfin la paix ?, préface de D. Lecourt, Le PUF, 3 vol., 5 000 p. Ouvrage couronné
Seuil. par l’Académie des Sciences.
• Gould, S. J. (2002) The structure of • Wright, R. (1995) L’Animal
evolutionary theory, Harvard University Moral, Michalon.
Press (anglophone). • Zimmer, K. (2001) Evolution :
• Grasset P. P. (1973) L'Évolution du the triumph of an idea, Harper Collins
vivant, matériaux pour une théorie (anglophone)
transformiste, Albin Michel.
• Grimoult, C. (2000) Histoire de
l'évolutionnisme contemporain en France
(1945-1995), Genève, Droz.
• Grimoult, C.(2001) L'évolution
biologique en France. Une révolution
scientifique, politique et culturelle, Genève,
Articles connexes [modifier]
• Adaptation • Ève • Mémétique
• Arbre mitochondriale • Panspermie
phylogénétique du • Évolution • Photo-guide
vivant convergente taxinomique du monde
• Biodiversité • Évolution animal
• Biologie de insulaire • Photo-guide
l'évolution • Exaptation taxinomique du monde
• Blessure • Fixisme végétal
narcissique • Gène égoïste • Prédéterminis
• Charles • Génération me
Darwin spontanée • Sélection
• Cladistique • Géographie naturelle
• Classification physique • Sélection de
phylogénétique du • Histoire de la parentèle
vivant pensée évolutionniste • Spéciation
• Classification • Historique des • Taxinomie
phylogénétique critiques des théories • Théorie du
• Coévolution de l'évolution handicap
• Conflit sexuel • Innovation • Théorie
• Créationnisme évolutive synthétique de
• Évolution • Institut l'évolution
Charles Darwin • Théorie
moléculaire international synergique de
• Évolution • Jean-Baptiste l'évolution
dirigée de Lamarck • William Donald
• Eukaryota Hamilton
• Équilibre

Liens externes [modifier]

• (fr) Évolution. De l'origine de la vie aux origines de l'homme, dossier SagaScience du
CNRS (France).
• Sélection de sites web sur la systématique et l’évolution dans le répertoire encyclopédique :
Les Signets de la Bibliothèque nationale de France
• Effervesciences (CINAPS Télévision) : Darwin aujourdhui (avec Guillaume Lecointre)
• (en) L'évolution résumée en 60 secondes

Évolution biologique

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et


Dérive génétique · Mutation génétique · Recombinaison

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant


History of evolutionary thought

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This article is about the history of evolutionary thought in biology. For the history of evolutionary
thought in the social sciences, see Sociocultural evolution. For the history of religious discussions, see
History of the creation-evolution controversy.
The Tree of Life as depicted by Ernst Haeckel in The Evolution of Man (1879) illustrates the 19th-
century view that evolution was a progressive process leading towards man.
Evolutionary thought, the conception that species change over time, has roots in antiquity, in the
ideas of the ancient Greeks, Romans, and Chinese as well as in medieval Islamic science. However, until the
18th century, Western biological thinking was dominated by essentialism, the belief that every species has
essential characteristics that are unalterable. This began to change during the Enlightenment when
evolutionary cosmology and the mechanical philosophy spread from the physical sciences to natural history.
Naturalists began to focus on the variability of species; the emergence of paleontology with the concept of
extinction further undermined the static view of nature. In the early 19th century, Jean-Baptiste Lamarck
proposed his theory of the transmutation of species, the first fully formed scientific theory of evolution.
In 1858, Charles Darwin and Alfred Russel Wallace published a new evolutionary theory that was
explained in detail in Darwin's On the Origin of Species (1859). Unlike Lamarck, Darwin proposed common
descent and a branching tree of life. The theory was based on the idea of natural selection, and it
synthesized a broad range of evidence from animal husbandry, biogeography, geology, morphology, and
The debate over Darwin's work led to the rapid acceptance of the general concept of evolution, but
the specific mechanism he proposed, natural selection, was not widely accepted until it was revived by
developments in biology that occurred during 1920s through the 1940s. Before that time most biologists
argued that other factors were responsible for evolution. Alternatives to natural selection suggested during
the eclipse of Darwinism included inheritance of acquired characteristics (neo-Lamarckism), an innate drive
for change (orthogenesis), and sudden large mutations (saltationism). The synthesis of natural selection with
Mendelian genetics during the 1920s and 1930s founded the new discipline of population genetics.
Throughout the 1930s and 1940s, population genetics became integrated with other biological fields,
resulting in a widely applicable theory of evolution that encompassed much of biology—the modern
evolutionary synthesis.
Following the establishment of evolutionary biology, studies of mutation and variation in natural
populations, combined with biogeography and systematics, led to sophisticated mathematical and causal
models of evolution. Paleontology and comparative anatomy allowed more detailed reconstructions of the
history of life. After the rise of molecular genetics in the 1950s, the field of molecular evolution developed,
based on protein sequences and immunological tests, and later incorporating RNA and DNA studies. The
gene-centered view of evolution rose to prominence in the 1960s, followed by the neutral theory of molecular
evolution, sparking debates over adaptationism, the units of selection, and the relative importance of genetic
drift versus natural selection. In the late 20th century, DNA sequencing led to molecular phylogenetics and
the reorganization of the tree of life into the three-domain system. In addition, the newly recognized factors of
symbiogenesis and horizontal gene transfer introduced yet more complexity into evolutionary history.
Part of the Biology series on

Mechanisms and processes

Genetic drift
Gene flow
Natural selection

Research and history

Evolutionary history of life
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Population genetics

Biology portal · v • d • e
• 1 Antiquity
• 1.1 Greeks
• 1.2 Chinese
• 1.3 Romans
• 1.4 Augustine of Hippo
• 2 Middle Ages
• 2.1 Islamic philosophy and the struggle for
• 2.2 Christian philosophy and the great chain of
• 2.3 Thomas Aquinas on creation and natural
• 3 Renaissance and Enlightenment
• 4 Early 19th century
• 4.1 Paleontology and geology
• 4.2 Transmutation of species
• 4.3 Anticipations of natural selection
• 4.4 Natural selection
• 5 1859–1930s: Darwin and his legacy
[edit] Antiquity
[edit] Greeks
Several ancient Greek philosophers discussed ideas that involved change in living organisms over
time. Anaximander (c.610–546 BC) proposed that the first animals lived in water and animals that live on land
were generated from them.[1] Empedocles (c. 490–430 BC) wrote of a non-supernatural origin for living
things,[2] suggesting that adaptation did not require an organizer or final cause. Aristotle summarized his
idea: "Wherever then all the parts came about just what they would have been if they had come to be for an
end, such things survived, being organized spontaneously in a fitting way; whereas those which grew
otherwise perished and continue to perish ..." although Aristotle himself rejected this view.[3]
Plato (left) and Aristotle (right), a detail of The School of Athens
Plato (c. 428–348 BC) was, in the words of biologist and historian Ernst Mayr, "the great antihero of
evolutionism",[4] as he established the philosophy of essentialism, which he called the Theory of Forms. This
theory holds that objects observed in the real world are only reflections of a limited number of essences
(eide). Variation merely results from an imperfect reflection of these constant essences. In his Timaeus, Plato
set forth the idea that the Demiurge had created the cosmos and everything in it because, being good, and
hence, "... free from jealousy, He desired that all things should be as like Himself as they could be". The
creator created all conceivable forms of life, since "... without them the universe will be incomplete, for it will
not contain every kind of animal which it ought to contain, if it is to be perfect". This "plenitude principle"—the
idea that all potential forms of life are essential to a perfect creation—greatly influenced Christian thought.[5]
Aristotle (384–322 BC), one of the most influential of the Greek philosophers, is the earliest natural
historian whose work has been preserved in any real detail. His writings on biology resulted from his
research into natural history on and around the isle of Lesbos, and have survived in the form of four books,
usually known by their Latin names, De anima (on the essence of life), Historia animalium (inquiries about
animals), De generatione animalium (reproduction), and De partibus animalium (anatomy). Aristotle's works
contain some remarkably astute observations and interpretations—along with sundry myths and mistakes—
reflecting the uneven state of knowledge during his time.[6] However, for Charles Singer, "Nothing is more
remarkable than [Aristotle's] efforts to [exhibit] the relationships of living things as a scala naturæ."[6] This
scala naturæ, described in Historia animalium, classified organisms in relation to a hierarchical "Ladder of
Life" or "Chain of Being", placing them according to their complexity of structure and function, with organisms
that showed greater vitality and ability to move described as "higher organisms".[5]

[edit] Chinese
Ideas on evolution were expressed by ancient Chinese thinkers such as Zhuangzi (Chuang Tzu), a
Taoist philosopher who lived around the 4th century BC. According to Joseph Needham, Taoism explicitly
denies the fixity of biological species and Taoist philosophers speculated that species had developed
differing attributes in response to differing environments.[7] Humans, nature and the heavens are seen as
existing in a state of "constant transformation" known as the Tao, in contrast with the more static view of
nature typical of Western thought.[8]
[edit] Romans
Titus Lucretius Carus (d. 50 BC), the Roman philosopher and atomist, wrote the poem On the Nature
of Things (De rerum natura), which provides the best surviving explanation of the ideas of the Greek
Epicurean philosophers. It describes the development of the cosmos, the Earth, living things, and human
society through purely naturalistic mechanisms, without any reference to supernatural involvement. On the
Nature of things would influence the cosmological and evolutionary speculations of philosophers and
scientists during and after the Renaissance.[9][10]

[edit] Augustine of Hippo

In line with earlier Greek thought, the 4th century bishop and theologian, St. Augustine of Hippo,
wrote that the creation story in Genesis should not be read too literally. In his book De Genesi ad litteram
("On The Literal Interpretation of Genesis"), he stated that in some cases new creatures may have come
about through the "decomposition" of earlier forms of life.[11] For Augustine, "plant, fowl and animal life are
not perfect ... but created in a state of potentiality", unlike what he considered the theologically perfect forms
of angels, the firmament and the human soul.[12] Augustine's idea 'that forms of life had been transformed
"slowly over time"' prompted Father Giuseppe Tanzella-Nitti, Professor of Theology at the Pontifical Santa
Croce University in Rome, to claim that Augustine had suggested a form of evolution.[13][14]

[edit] Middle Ages

[edit] Islamic philosophy and the struggle for existence
See also: Early Islamic philosophy and Science in medieval Islam
Although Greek and Roman evolutionary ideas died out in Europe after the fall of the Roman Empire,
they were not lost to Islamic philosophers and scientists. In the Islamic Golden Age of the 8th to the 13th
centuries, philosophers explored ideas about natural history. These ideas included transmutation from non-
living to living: "from mineral to plant, from plant to animal, and from animal to man".[15]
The first Muslim biologist and philosopher to speculate in detail about natural history was the Afro-
Arab writer al-Jahiz in the 9th century. In the Book of Animals, he considered the effects of the environment
on an animal's chances for survival, and described the struggle for existence.[16] Al-Jahiz also wrote
descriptions of food chains.[17] Al-Jahiz speculated on the influence of the environment on animals and
considered the effects of the environment on the likelihood of an animal to survive. For example, Al-Jahiz's
wrote in his Book of Animals:
All animals, in short, can not exist without food, neither can the hunting animal escape being
hunted in his turn. Every weak animal devours those weaker than itself. Strong animals cannot
escape being devoured by other animals stronger than they.[16]
[edit] Christian philosophy and the great chain of being
Drawing of the great chain of being from Rhetorica Christiana (1579) by Diego Valades
During the Early Middle Ages, Greek classical learning was all but lost to the West. However, contact
with the Islamic world, where Greek manuscripts were preserved and expanded, soon led to a massive spate
of Latin translations in the 12th century. Europeans were re-introduced to the works of Plato and Aristotle, as
well as to Islamic thought. Christian thinkers of the scholastic school, in particular Abelard and Thomas
Aquinas, combined Aristotelian classification with Plato's ideas of the goodness of God, and of all potential
life forms being present in a perfect creation, to organize all inanimate, animate, and spiritual beings into a
huge interconnected system: the scala naturæ, or great chain of being.[5][18]
Within this system, everything that existed could be placed in order, from "lowest" to "highest", with
Hell at the bottom and God at the top—below God, an angelic hierarchy marked by the orbits of the planets,
mankind in an intermediate position, and worms the lowest of the animals. As the universe was ultimately
perfect, the great chain was also perfect. There were no empty links in the chain, and no link was
represented by more than one species. Therefore no species could ever move from one position to another.
Thus, in this Christianized version of Plato's perfect universe, species could never change, but remained
forever fixed, in accordance with the text of Genesis. For humans to forget their position was seen as sinful,
whether they behaved like lower animals or aspired to a higher station than was given them by their Creator.
Creatures on adjacent steps were expected to closely resemble each other, an idea expressed in the
saying: natura non facit saltum ("nature does not make leaps").[5] This basic concept of the great chain of
being greatly influenced the thinking of Western civilization for centuries (and still has an influence today). It
formed a part of the argument from design presented by natural theology. As a classification system, it
became the major organizing principle and foundation of the emerging science of biology in the 17th and
18th centuries.[5]
[edit] Thomas Aquinas on creation and natural processes
While the development of the great chain of being and the argument from design by Christian
theologians contributed to the view that the natural world fit into an unchanging designed hierarchy, some
theologians were more open to the possibility that the world might have developed through natural
processes. Thomas Aquinas went even farther than Augustine of Hippo in arguing that scriptural texts like
Genesis should not be interpreted in a literal way that conflicted with or constrained what natural
philosophers learned about the workings of the natural world. He felt that the autonomy of nature was a sign
of God's goodness and that there was no conflict between the concept of a divinely created universe, and the
idea that the universe may have evolved over time through natural mechanisms.[19] However, Aquinas
disputed the views of those like the ancient Greek philosopher Empodocles who held that such natural
processes showed that the universe could have developed without an underlying purpose. Rather holding
Hence, it is clear that nature is nothing but a certain kind of art, i.e., the divine art, impressed
upon things, by which these things are moved to a determinate end. It is as if the shipbuilder
were able to give to timbers that by which they would move themselves to take the form of a

[edit] Renaissance and Enlightenment

Main article: Evolutionary ideas of the Renaissance and Enlightenment
Pierre Belon compared the skeletons of birds and humans in his Book of Birds (1555).
In the first half of the 17th century, René Descartes's mechanical philosophy encouraged the use of
the metaphor of the universe as a machine, a concept that would come to characterise the scientific
revolution.[21] Between 1650 and 1800, some evolutionist theories supported the view that the universe,
including life on Earth, had developed mechanically, entirely without divine guidance. In contrast, most
contemporary theories of evolution, such of those of Gottfried Leibniz and J. G. Herder, held that evolution
was a fundamentally spiritual process.[22] In 1751, Pierre Louis Maupertuis veered toward more materialist
ground. He wrote of natural modifications occurring during reproduction and accumulating over the course of
many generations, producing races and even new species, a description that anticipated in general terms the
concept of natural selection.[23]
Later in the 18th century, the French philosopher G. L. L. Buffon, one of the leading 18th century
naturalists, suggested that what most people referred to as species were really just well-marked varieties,
modified from an original form by environmental factors. For example, he believed that lions, tigers, leopards
and house cats might all have a common ancestor. He further speculated that the 200 or so species of
mammals then known might have descended from as few as 38 original animal forms. Buffon's evolutionary
ideas were limited; he believed each of the original forms had arisen through spontaneous generation and
that each was shaped by "internal moulds" that limited the amount of change. Buffon's works, Natural History
and The Epochs of Nature, containing well developed theories about a completely materialistic origin for the
Earth and his ideas questioning the fixity of species, were extremely influential.[24][25]
Between 1767 and 1792, James Burnett, Lord Monboddo included in his writings not only the
concept that man had descended from primates, but also that, in response to the environment, creatures had
found methods of transforming their characteristics over long time intervals.[26] Charles Darwin's
grandfather, Erasmus Darwin, published Zoönomia in 1796, which suggested that "all warm-blooded animals
have arisen from one living filament".[27] In his 1802 poem Temple of Nature, he described the rise of life
from minute organisms living in mud to all of its modern diversity.[28]
[edit] Early 19th century
Diagram of the geologic timescale from an 1861 book by Richard Owen showing the appearance of
major animal types

[edit] Paleontology and geology

See also: History of paleontology
In 1796, Georges Cuvier published his findings on the differences between living elephants and
those found in the fossil record. His analysis demonstrated that mammoths and mastodons were distinct
species different from any living animal, effectively ending a long-running debate over whether the extinction
of a species was possible.[29] In 1788, James Hutton described gradual geological processes operating
continuously over deep time.[30] In the 1790s William Smith began the process of ordering rock strata by
examining fossils in the layers while he worked on his geologic map of England. Independently, in 1811,
Georges Cuvier and Alexandre Brongniart published an influential study of the geologic history of the region
around Paris, based on the stratigraphic succession of rock layers. These works helped establish the
antiquity of the Earth.[31] Cuvier advocated catastrophism to explain the patterns of extinction and faunal
succession revealed by the fossil record.
Knowledge of the fossil record continued to advance rapidly during the first few decades of the 19th
century. By the 1840s, the outlines of the geologic timescale were becoming clear, and in 1841 John Phillips
named three major eras, based on the predominant fauna of each: the Paleozoic, dominated by marine
invertebrates and fish, the Mesozoic, the age of reptiles, and the current Cenozoic age of mammals. This
progressive picture of the history of life was accepted even by conservative English geologists like Adam
Sedgwick and William Buckland; however, like Cuvier, they attributed the progression to repeated
catastrophic episodes of extinction followed by new episodes of creation.[32] Unlike Cuvier, Buckland and
some other advocates of natural theology among British geologists made efforts to explicitly link the last
catastrophic episode proposed by Cuvier to the biblical flood.[33][34]
From 1830 to 1833, Charles Lyell published his multi-volume work Principles of Geology, which,
building on Hutton's ideas, advocated a uniformitarian alternative to the catastrophic theory of geology. Lyell
claimed that, rather than being the products of cataclysmic (and possibly supernatural) events, the geologic
features of the Earth are better explained as the result of the same gradual geologic forces observable in the
present day—but acting over immensely long periods of time. Although Lyell opposed evolutionary ideas
(even questioning the consensus that the fossil record demonstrates a true progression), his concept that the
Earth was shaped by forces working gradually over an extended period, and the immense age of the Earth
assumed by his theories, would strongly influence future evolutionary thinkers such as Charles Darwin.[35]

[edit] Transmutation of species

Main article: Transmutation of species
Diagram from Vestiges of the Natural History of Creation (1844) by Robert Chambers shows a model
of development where fish (F), reptiles (R), and birds (B) represent branches from a path leading to
mammals (M).
Jean-Baptiste Lamarck proposed, in his Philosophie Zoologique of 1809, a theory of the
transmutation of species. Lamarck did not believe that all living things shared a common ancestor but rather
that simple forms of life were created continuously by spontaneous generation. He also believed that an
innate life force drove species to become more complex over time, advancing up a linear ladder of
complexity that was related to the great chain of being. Lamarck recognized that species were adapted to
their environment. He explained this by saying that the same innate force driving increasing complexity
caused the organs of an animal (or a plant) to change based on the use or disuse of those organs, just as
muscles are affected by exercise. He argued that these changes would be inherited by the next generation
and produce slow adaptation to the environment. It was this secondary mechanism of adaptation through the
inheritance of acquired characteristics that would become known as Lamarckism and would influence
discussions of evolution into the 20th century.[36][37]
A radical British school of comparative anatomy that included the anatomist Robert Grant was closely
in touch with Lamarck's French school of Transformationism. One of the French scientists who influenced
Grant was the anatomist Étienne Geoffroy Saint-Hilaire, whose ideas on the unity of various animal body
plans and the homology of certain anatomical structures would be widely influential and lead to intense
debate with his colleague Georges Cuvier. Grant became an authority on the anatomy and reproduction of
marine invertebrates. He developed Lamarck's and Erasmus Darwin's ideas of transmutation and
evolutionism, and investigated homology, even proposing that plants and animals had a common
evolutionary starting point. As a young student Charles Darwin joined Grant in investigations of the life cycle
of marine animals. In 1826 an anonymous paper, probably written by Robert Jameson, praised Lamarck for
explaining how higher animals had "evolved" from the simplest worms; this was the first use of the word
"evolved" in a modern sense.[38][39]
In 1844, the Scottish publisher Robert Chambers anonymously published an extremely controversial
but widely read book entitled Vestiges of the Natural History of Creation. This book proposed an evolutionary
scenario for the origins of the Solar System and life on Earth. It claimed that the fossil record showed a
progressive ascent of animals with current animals being branches off a main line that leads progressively to
humanity. It implied that the transmutations lead to the unfolding of a preordained plan that had been woven
into the laws that governed the universe. In this sense it was less completely materialistic than the ideas of
radicals like Robert Grant, but its implication that humans were only the last step in the ascent of animal life
incensed many conservative thinkers. The high profile of the public debate over Vestiges, with its depiction of
evolution as a progressive process, would greatly influence the perception of Darwin's theory a decade later.
Ideas about the transmutation of species were associated with the radical materialism of the
Enlightenment and were attacked by more conservative thinkers. Georges Cuvier attacked the ideas of
Lamarck and Geoffroy Saint-Hilaire, agreeing with Aristotle that species were immutable. Cuvier believed
that the individual parts of an animal were too closely correlated with one another to allow for one part of the
anatomy to change in isolation from the others, and argued that the fossil record showed patterns of
catastrophic extinctions followed by re-population, rather than gradual change over time. He also noted that
drawings of animals and animal mummies from Egypt, which were thousands of years old, showed no signs
of change when compared with modern animals. The strength of Cuvier's arguments and his scientific
reputation helped keep transmutational ideas out of the mainstream for decades.[42]
This 1847 diagram by Richard Owen shows his conceptual archetype for all vertebrates.
In Britain the philosophy of natural theology remained influential. William Paley's 1802 book Natural
Theology with its famous watchmaker analogy had been written at least in part as a response to the
transmutational ideas of Erasmus Darwin.[43] Geologists influenced by natural theology, such as Buckland
and Sedgwick, made a regular practice of attacking the evolutionary ideas of Lamarck, Grant, and The
Vestiges of the Natural History of Creation.[44][45] Although the geologist Charles Lyell opposed scriptural
geology, he also believed in the immutability of species, and in his Principles of Geology (1830–1833), he
criticized Lamarck's theories of development.[35] Idealists such as Louis Agassiz and Richard Owen believed
that each species was fixed and unchangeable because it represented an idea in the mind of the creator.
They believed that relationships between species could be discerned from developmental patterns in
embryology, as well as in the fossil record, but that these relationships represented an underlying pattern of
divine thought, with progressive creation leading to increasing complexity and culminating in humanity. Owen
developed the idea of "archetypes" in the Divine mind that would produce a sequence of species related by
anatomical homologies, such as vertebrate limbs. Owen led a public campaign that successfully
marginalized Robert Grant in the scientific community. Darwin would make good use of the homologies
analyzed by Owen in his own theory, but the harsh treatment of Grant, and the controversy surrounding
Vestiges, showed him the need to ensure that his own ideas were scientifically sound.[39][46][47]

[edit] Anticipations of natural selection

Several writers anticipated aspects of Darwin's theory, and in the third edition of On the Origin of
Species published in 1861 Darwin named those he knew about in an introductory appendix, An Historical
Sketch of the Recent Progress of Opinion on the Origin of Species , which he expanded in later editions.[48]
In 1813, William Charles Wells read before the Royal Society essays assuming that there had been
evolution of humans, and recognising the principle of natural selection. Charles Darwin and Alfred Russel
Wallace were unaware of this work when they jointly published the theory in 1858, but Darwin later
acknowledged that Wells had recognised the principle before them, writing that the paper "An Account of a
White Female, part of whose Skin resembles that of a Negro" was published in 1818, and "he distinctly
recognises the principle of natural selection, and this is the first recognition which has been indicated; but he
applies it only to the races of man, and to certain characters alone."[49] When Darwin was developing his
theory, he was influenced by Augustin de Candolle's natural system of classification, which laid emphasis on
the war between competing species.[50][51]
Patrick Matthew wrote in the obscure book Naval Timber & Arboriculture (1831) of "continual
balancing of life to circumstance. ... [The] progeny of the same parents, under great differences of
circumstance, might, in several generations, even become distinct species, incapable of co-
reproduction."[52] Charles Darwin discovered this work after the initial publication of the Origin. In the brief
historical sketch that Darwin included in the 3rd edition he says "Unfortunately the view was given by Mr.
Matthew very briefly in an Appendix to a work on a different subject ... He clearly saw, however, the full force
of the principle of natural selection."[53]
It is possible to look through the history of biology from the ancient Greeks onwards and discover
anticipations of almost all of Darwin's key ideas. However, as historian of science Peter J. Bowler says,
"Through a combination of bold theorizing and comprehensive evaluation, Darwin came up with a concept of
evolution that was unique for the time." Bowler goes on to say that simple priority alone is not enough to
secure a place in the history of science; someone has to develop an idea and convince others of its
importance to have a real impact.[54]
T. H. Huxley said in his essay on the reception of the Origin of Species:
The suggestion that new species may result from the selective action of external conditions
upon the variations from their specific type which individuals present and which we call
spontaneous because we are ignorant of their causation is as wholly unknown to the historian of
scientific ideas as it was to biological specialists before 1858. But that suggestion is the central
idea of the Origin of Species, and contains the quintessence of Darwinism.[55]
Darwin's first sketch of an evolutionary tree from his First Notebook on Transmutation of Species

[edit] Natural selection

Main articles: Inception of Darwin's theory, Development of Darwin's theory, and Publication of
Darwin's theory
The biogeographical patterns Charles Darwin observed in places such as the Galapagos islands
during the voyage of the Beagle caused him to doubt the fixity of species, and in 1837 Darwin started the first
of a series of secret notebooks on transmutation. Darwin's observations led him to view transmutation as a
process of divergence and branching, rather than the ladder-like progression envisioned by Lamarck and
others. In 1838 he read the new 6th edition of An Essay on the Principle of Population, written in the late
1700s by Thomas Malthus. Malthus' idea of population growth leading to a struggle for survival combined
with Darwin's knowledge on how breeders selected traits, led to the inception of Darwin's theory of natural
selection. Darwin did not publish his ideas on evolution for 20 years. However he did share them with certain
other naturalists and friends, starting with Joseph Hooker, with whom he discussed his unpublished 1844
essay on natural selection. During this period he used the time he could spare from his other scientific work
to slowly refine his ideas and, aware of the intense controversy around transmutation, amass evidence to
support them. In September 1854 he began full time work on writing his book on natural selection.[47][56][57]
Unlike Darwin, Alfred Russel Wallace, influenced by the book Vestiges of the Natural History of
Creation, already suspected that transmutation of species occurred when he began his career as a naturalist.
By 1855 his biogeographical observations during his field work in South America and the Malay Archipelago
made him confident enough in a branching pattern of evolution to publish a paper stating that every species
originated in close proximity to an already existing closely allied species. Like Darwin, it was Wallace's
consideration of how the ideas of Malthus might apply to animal populations that led him to conclusions very
similar to those reached by Darwin about the role of natural selection. In February 1858 Wallace, unaware of
Darwin's unpublished ideas, composed his thoughts into an essay and mailed them to Darwin, asking for his
opinion. The result was the joint publication in July of an extract from Darwin's 1844 essay along with
Wallace's letter. Darwin also began work on a short abstract summarising his theory, which he would publish
in 1859 as On the Origin of Species.[58]
Diagram by O.C. Marsh of the evolution of horse feet and teeth over time as reproduced in T.H
Huxley's 1876 book Professor Huxley in America
[edit] 1859–1930s: Darwin and his legacy
See also: Reaction to Darwin's theory
By the 1850s whether or not species evolved was a subject of intense debate, with prominent
scientists arguing both sides of the issue.[59] However, it was the publication of Charles Darwin's On the
Origin of Species (1859) that fundamentally transformed the discussion over biological origins.[60] Darwin
argued that his branching version of evolution explained a wealth of facts in biogeography, anatomy,
embryology, and other fields of biology. He also provided the first cogent mechanism by which evolutionary
change could persist: his theory of natural selection.[61]
One of the first and most important naturalists to be convinced by Origin of the reality of evolution
was the British anatomist Thomas Henry Huxley. Huxley recognized that unlike the earlier transmutational
ideas of Lamarck and Vestiges, Darwin's theory provided a mechanism for evolution without supernatural
involvement, even if Huxley himself was not completely convinced that natural selection was the key
evolutionary mechanism. Huxley would make advocacy of evolution a cornerstone of the program of the X
Club to reform and professionalise science by displacing natural theology with naturalism and to end the
domination of British natural science by the clergy. By the early 1870s in English-speaking countries, thanks
partly to these efforts, evolution had become the mainstream scientific explanation for the origin of species.
[61] In his campaign for public and scientific acceptance of Darwin's theory, Huxley made extensive use of
new evidence for evolution from paleontology. This included evidence that birds had evolved from reptiles,
including the discovery of Archaeopteryx in Europe, and a number of fossils of primitive birds with teeth found
in North America. Another important line of evidence was the finding of fossils that helped trace the evolution
of the horse from its small five-toed ancestors.[62] However, acceptance of evolution among scientists in
non-English speaking nations such as France, and the countries of southern Europe and Latin America was
slower. An exception to this was Germany, where both August Weismann and Ernst Haeckel championed
this idea: Haeckel used evolution to challenge the established tradition of metaphysical idealism in German
biology, much as Huxley used it to challenge natural theology in Britain.[63] Haeckel and other German
scientists would take the lead in launching an ambitious programme to reconstruct the evolutionary history of
life based on morphology and embryology.[64]
Darwin's theory succeeded in profoundly altering scientific opinion regarding the development of life
and in producing a small philosophical revolution.[65] However, this theory could not explain several critical
components of the evolutionary process. Specifically, Darwin was unable to explain the source of variation in
traits within a species, and could not identify a mechanism that could pass traits faithfully from one
generation to the next. Darwin's hypothesis of pangenesis, while relying in part on the inheritance of acquired
characteristics, proved to be useful for statistical models of evolution that were developed by his cousin
Francis Galton and the "biometric" school of evolutionary thought. However, this idea proved to be of little
use to other biologists.[66]
[edit] Application to humans

This illustration was the frontispiece of Thomas Henry Huxley's book Evidence as to Man's Place in
Nature (1863).
Charles Darwin was aware of the severe reaction in some parts of the scientific community against
the suggestion made in Vestiges of the Natural History of Creation that humans had arisen from animals by a
process of transmutation. Therefore he almost completely ignored the topic of human evolution in The Origin
of Species. Despite this precaution, the issue featured prominently in the debate that followed the book's
publication. For most of the first half of the 19th century, the scientific community believed that, although
geology had shown that the Earth and life were very old, human beings had appeared suddenly just a few
thousand years before the present. However, a series of archaeological discoveries in the 1840s and 1850s
showed stone tools associated with the remains of extinct animals. By the early 1860s, as summarized in
Charles Lyell's 1863 book Geological Evidences of the Antiquity of Man , it had become widely accepted that
humans had existed during a prehistoric period – which stretched many thousands of years before the start of
written history. This view of human history was more compatible with an evolutionary origin for humanity than
was the older view. On the other hand, at that time there was no fossil evidence to demonstrate human
evolution. The only human fossils found before the discovery of Java man in the 1890s were either of
anatomically modern humans or of Neanderthals that were too close, especially in the critical characteristic
of cranial capacity, to modern humans for them to be convincing intermediates between humans and other
Therefore the debate that immediately followed the publication of The Origin of Species centered on
the similarities and differences between humans and modern apes. Carolus Linnaeus had been criticised in
the 18th century for grouping humans and apes together as primates in his ground breaking classification
system.[68] Richard Owen vigorously defended the classification suggested by Cuvier and Johann Friedrich
Blumenbach that placed humans in a separate order from any of the other mammals, which by the early 19th
century had become the orthodox view. On the other hand, Thomas Henry Huxley sought to demonstrate a
close anatomical relationship between humans and apes. In one famous incident, Huxley showed that Owen
was mistaken in claiming that the brains of gorillas lacked a structure present in human brains. Huxley
summarized his argument in his highly influential 1863 book Evidence as to Man's Place in Nature. Another
viewpoint was advocated by Charles Lyell and Alfred Russel Wallace. They agreed that humans shared a
common ancestor with apes, but questioned whether any purely materialistic mechanism could account for
all the differences between humans and apes, especially some aspects of the human mind.[67]
In 1871, Darwin published The Descent of Man, and Selection in Relation to Sex , which contained
his views on human evolution. Darwin argued that the differences between the human mind and the minds of
the higher animals were a matter of degree rather than of kind. For example, he viewed morality as a natural
outgrowth of instincts that were beneficial to animals living in social groups. He argued that all the differences
between humans and apes were explained by a combination of the selective pressures that came from our
ancestors moving from the trees to the plains, and sexual selection. The debate over human origins, and
over the degree of human uniqueness continued well into the 20th century.[67]

[edit] Alternatives to natural selection

Main article: The eclipse of Darwinism
This photo from Henry Fairfield Osborn's 1918 book Origin and Evolution of Life shows models
depicting the evolution of Titanothere horns over time, which Osborn claimed was an example of an
orthogenic trend in evolution.
The concept of evolution was widely accepted in scientific circles within a few years of the publication
of Origin, but the acceptance of natural selection as its driving mechanism was much less widespread. The
four major alternatives to natural selection in the late 19th century were theistic evolution, neo-Lamarckism,
orthogenesis, and saltationism. Theistic evolution (a term promoted by Darwin's greatest American advocate
Asa Gray) was the idea that God intervened in the process of evolution to guide it in such a way that the
living world could still be considered to be designed. However, this idea gradually fell out of favor among
scientists, as they became more and more committed to the idea of methodological naturalism and came to
believe that direct appeals to supernatural involvement were scientifically unproductive. By 1900, theistic
evolution had largely disappeared from professional scientific discussions, although it retained a strong
popular following.[69][70]
In the late 19th century, the term neo-Lamarckism came to be associated with the position of
naturalists who viewed the inheritance of acquired characteristics as the most important evolutionary
mechanism. Advocates of this position included the British writer and Darwin critic Samuel Butler, the
German biologist Ernst Haeckel, and the American paleontologist Edward Drinker Cope. They considered
Lamarckism to be philosophically superior to Darwin's idea of selection acting on random variation. Cope
looked for, and thought he found, patterns of linear progression in the fossil record. Inheritance of acquired
characteristics was part of Haeckel's recapitulation theory of evolution, which held that the embryological
development of an organism repeats its evolutionary history.[69][70] Critics of neo-Lamarckism, such as the
German biologist August Weismann and Alfred Russel Wallace, pointed out that no one had ever produced
solid evidence for the inheritance of acquired characteristics. Despite these criticisms, neo-Lamarckism
remained the most popular alternative to natural selection at the end of the 19th century, and would remain
the position of some naturalists well into the 20th century.[69][70]
Orthogenesis was the hypothesis that life has an innate tendency to change, in a unilinear fashion,
towards ever-greater perfection. It had a significant following in the 19th century, and its proponents included
the Russian biologist Leo Berg and the American paleontologist Henry Fairfield Osborn. Orthogenesis was
popular among some paleontologists, who believed that the fossil record showed a gradual and constant
unidirectional change. Saltationism was the idea that new species arise as a result of large mutations. It was
seen as a much faster alternative to the Darwinian concept of a gradual process of small random variations
being acted on by natural selection, and was popular with early geneticists such as Hugo de Vries, William
Bateson, and early in his career, T. H. Morgan. It became the basis of the mutation theory of evolution.[69]
Diagram from T.H. Morgan's 1919 book The Physical Basis of Heredity, showing the sex-linked
inheritance of the white-eyed mutation in Drosophila melanogaster

[edit] Mendelian genetics, biometrics, and mutation

The so-called rediscovery of Gregor Mendel's laws of inheritance in 1900 ignited a fierce debate
between two camps of biologists. In one camp were the Mendelians, who were focused on discrete variations
and the laws of inheritance. They were led by William Bateson (who coined the word genetics) and Hugo de
Vries (who coined the word mutation). Their opponents were the biometricians, who were interested in the
continuous variation of characteristics within populations. Their leaders, Karl Pearson and Walter Frank
Raphael Weldon, followed in the tradition of Francis Galton, who had focused on measurement and statistical
analysis of variation within a population. The biometricians rejected Mendelian genetics on the basis that
discrete units of heredity, such as genes, could not explain the continuous range of variation seen in real
populations. Weldon's work with crabs and snails provided evidence that selection pressure from the
environment could shift the range of variation in wild populations, but the Mendelians maintained that the
variations measured by biometricians were too insignificant to account for the evolution of new species.[71]
When T. H. Morgan began experimenting with breeding the fruit fly Drosophila melanogaster, he was
a saltationist who hoped to demonstrate that a new species could be created in the lab by mutation alone.
Instead, the work at his lab between 1910 and 1915 reconfirmed Mendelian genetics and provided solid
experimental evidence linking it to chromosomal inheritance. His work also demonstrated that most
mutations had relatively small effects, such as a change in eye color, and that rather than creating a new
species in a single step, mutations served to increase variation within the existing population.[71][72]
[edit] 1920s–1940s
See also: Modern evolutionary synthesis

Biston betularia f. typica is the white-bodied form of the peppered moth.

Biston betularia f. carbonaria is the black-bodied form of the peppered moth.

[edit] Population genetics
The Mendelian and biometrician models were eventually reconciled with the development of
population genetics. A key step was the work of the British biologist and statistician R.A. Fisher. In a series of
papers starting in 1918 and culminating in his 1930 book The Genetical Theory of Natural Selection, Fisher
showed that the continuous variation measured by the biometricians could be produced by the combined
action of many discrete genes, and that natural selection could change gene frequencies in a population,
resulting in evolution. In a series of papers beginning in 1924, another British geneticist, J.B.S. Haldane,
applied statistical analysis to real-world examples of natural selection, such as the evolution of industrial
melanism in peppered moths, and showed that natural selection worked at an even faster rate than Fisher
The American biologist Sewall Wright, who had a background in animal breeding experiments,
focused on combinations of interacting genes, and the effects of inbreeding on small, relatively isolated
populations that exhibited genetic drift. In 1932, Wright introduced the concept of an adaptive landscape and
argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive
peak, allowing natural selection to drive it towards different adaptive peaks. The work of Fisher, Haldane and
Wright founded the discipline of population genetics. This integrated natural selection with Mendelian
genetics, which was the critical first step in developing a unified theory of how evolution worked.[73][74]

[edit] Modern evolutionary synthesis

In the first few decades of the 20th century, most field naturalists continued to believe that
Lamarckian and orthogenic mechanisms of evolution provided the best explanation for the complexity they
observed in the living world. But as the field of genetics continued to develop, those views became less
tenable.[75] Theodosius Dobzhansky, a postdoctoral worker in T. H. Morgan's lab, had been influenced by
the work on genetic diversity by Russian geneticists such as Sergei Chetverikov. He helped to bridge the
divide between the foundations of microevolution developed by the population geneticists and the patterns of
macroevolution observed by field biologists, with his 1937 book Genetics and the Origin of Species.
Dobzhansky examined the genetic diversity of wild populations and showed that, contrary to the assumptions
of the population geneticists, these populations had large amounts of genetic diversity, with marked
differences between sub-populations. The book also took the highly mathematical work of the population
geneticists and put it into a more accessible form. In Great Britain E.B. Ford, the pioneer of ecological
genetics, continued throughout the 1930s and 1940s to demonstrate the power of selection due to ecological
factors including the ability to maintain genetic diversity through genetic polymorphisms such as human
blood types. Ford's work would contribute to a shift in emphasis during the course of the modern synthesis
towards natural selection over genetic drift.[73][74][76][77]
Evolutionary biologist Ernst Mayr was influenced by the work of the German biologist Bernhard
Rensch showing the influence of local environmental factors on the geographic distribution of sub-species
and closely related species. Mayr followed up on Dobzhansky's work with the 1942 book Systematics and
the Origin of Species, which emphasized the importance of allopatric speciation in the formation of new
species. This form of speciation occurs when the geographical isolation of a sub-population is followed by the
development of mechanisms for reproductive isolation. Mayr also formulated the biological species concept
that defined a species as a group of interbreeding or potentially interbreeding populations that were
reproductively isolated from all other populations.[73][74][78]
In the 1944 book Tempo and Mode in Evolution, George Gaylord Simpson showed that the fossil
record was consistent with the irregular non-directional pattern predicted by the developing evolutionary
synthesis, and that the linear trends that earlier paleontologists had claimed supported orthogenesis and
neo-Lamarckism did not hold up to closer examination. In 1950, G. Ledyard Stebbins published Variation
and Evolution in Plants, which helped to integrate botany into the synthesis. The emerging cross-disciplinary
consensus on the workings of evolution would be known as the modern evolutionary synthesis. It received its
name from the book Evolution: The Modern Synthesis by Julian Huxley.[73][74]
The evolutionary synthesis provided a conceptual core—in particular, natural selection and Mendelian
population genetics—that tied together many, but not all, biological disciplines. It helped establish the
legitimacy of evolutionary biology, a primarily historical science, in a scientific climate that favored
experimental methods over historical ones.[79] The synthesis also resulted in a considerable narrowing of
the range of mainstream evolutionary thought (what Stephen Jay Gould called the "hardening of the
synthesis"): by the 1950s, natural selection acting on genetic variation was virtually the only acceptable
mechanism of evolutionary change (panselectionism), and macroevolution was simply considered the result
of extensive microevolution.[80][81]

[edit] 1940s–1960s: Molecular biology and evolution

Main article: History of molecular evolution
The middle decades of the 20th century saw the rise of molecular biology, and with it an
understanding of the chemical nature of genes as sequences of DNA and their relationship, through the
genetic code, to protein sequences. At the same time, increasingly powerful techniques for analyzing
proteins, such as protein electrophoresis and sequencing, brought biochemical phenomena into realm of the
synthetic theory of evolution. In the early 1960s, biochemists Linus Pauling and Emile Zuckerkandl proposed
the molecular clock hypothesis: that sequence differences between homologous proteins could be used to
calculate the time since two species diverged. By 1969, Motoo Kimura and others provided a theoretical
basis for the molecular clock, arguing that—at the molecular level at least—most genetic mutations are neither
harmful nor helpful and that genetic drift, rather than natural selection, is responsible for a large portion of
genetic change: the neutral theory of molecular evolution.[82] Studies of protein differences within species
also brought molecular data to bear on population genetics by providing estimates of the level of
heterozygosity in natural populations.[83]
From the early 1960s, molecular biology was increasingly seen as a threat to the traditional core of
evolutionary biology. Established evolutionary biologists—particularly Ernst Mayr, Theodosius Dobzhansky
and G. G. Simpson, three of the architects of the modern synthesis—were extremely skeptical of molecular
approaches, especially when it came to the connection (or lack thereof) to natural selection. The molecular
clock hypothesis and the neutral theory were particularly controversial, spawning the neutralist-selectionist
debate over the relative importance of drift and selection, which continued into the 1980s without a clear

[edit] Late 20th century

[edit] Gene-centered view
In the mid-1960s, George C. Williams strongly critiqued explanations of adaptations worded in terms
of "survival of the species" (group selection arguments). Such explanations were largely replaced by a gene-
centered view of evolution, epitomized by the kin selection arguments of W. D. Hamilton, George R. Price
and John Maynard Smith.[86] This viewpoint would be summarized and popularized in the influential 1976
book The Selfish Gene by Richard Dawkins.[87] Models of the period showed that group selection was
severely limited in its strength; though newer models do admit the possibility of significant multi-level
In 1973, Leigh Van Valen proposed the term "Red Queen", which he took from Through the Looking-
Glass by Lewis Carroll, to describe a scenario where a species involved in one or more evolutionary arms
races would have to constantly change just to keep pace with the species with which it was co-e