In biology, adaptation has three related meanings. Firstly it is the
dynamic evolutionary process that fits organisms to their environment,
enhancing their evolutionary fitness. Secondly, it is a state reached
by the population during that process. Thirdly, it is a phenotypic or
adaptive trait, with a functional role in each individual organism,
that is maintained and has been evolved by natural selection.
Organisms face a succession of environmental challenges as they grow,
and show adaptive plasticity as traits develop in response to the
imposed conditions. This gives them resilience to varying
2 General principles
2.1 What adaptation is
2.2 What adaptation is not
2.3 Adaptedness and fitness
2.4 Genetic basis
3.1 Changes in habitat
3.2 Genetic change
4 Shifts in function
4.2 Co-option of existing traits: exaptation
5 Non-adaptive traits
Extinction and coextinction
7 Philosophical issues
8 See also
Main article: History of evolutionary thought
Adaptation is an observable fact of life accepted by philosophers and
natural historians from ancient times, independently of their views on
evolution, but their explanations differed.
Empedocles did not believe
that adaptation required a final cause (~ purpose), but "came about
naturally, since such things survived."
Aristotle did believe in final
causes, but assumed that species were fixed.
In natural theology, adaptation was interpreted as the work of a deity
and as evidence for the existence of God.
William Paley believed
that organisms were perfectly adapted to the lives they led, an
argument that shadowed Gottfried Wilhelm Leibniz, who had argued that
God had brought about "the best of all possible worlds." Voltaire's
Dr. Pangloss is a parody of this optimistic idea, and David Hume
also argued against design. The Bridgewater Treatises are a product
of natural theology, though some of the authors managed to present
their work in a fairly neutral manner. The series was lampooned by
Robert Knox, who held quasi-evolutionary views, as the Bilgewater
Charles Darwin broke with the tradition by emphasising the
flaws and limitations which occurred in the animal and plant
Jean-Baptiste Lamarck proposed a tendency for organisms to become more
complex, moving up a ladder of progress, plus "the influence of
circumstances," usually expressed as use and disuse. This second,
subsidiary element of his theory is what is now called Lamarckism, a
proto-evolutionary hypothesis of the inheritance of acquired
characteristics, intended to explain adaptations by natural means.
Other natural historians, such as Buffon, accepted adaptation, and
some also accepted evolution, without voicing their opinions as to the
mechanism. This illustrates the real merit of Darwin and Alfred Russel
Wallace, and secondary figures such as Henry Walter Bates, for putting
forward a mechanism whose significance had only been glimpsed
previously. A century later, experimental field studies and breeding
experiments by people such as
E. B. Ford
E. B. Ford and Theodosius Dobzhansky
produced evidence that natural selection was not only the 'engine'
behind adaptation, but was a much stronger force than had previously
The significance of an adaptation can only be understood in relation
to the total biology of the species.
— Julian Huxley, Evolution: The Modern Synthesis
What adaptation is
Adaptation is primarily a process rather than a physical form or part
of a body. An internal parasite (such as a liver fluke) can
illustrate the distinction: such a parasite may have a very simple
bodily structure, but nevertheless the organism is highly adapted to
its specific environment. From this we see that adaptation is not just
a matter of visible traits: in such parasites critical adaptations
take place in the life cycle, which is often quite complex.
However, as a practical term, "adaptation" often refers to a product:
those features of a species which result from the process. Many
aspects of an animal or plant can be correctly called adaptations,
though there are always some features whose function remains in doubt.
By using the term adaptation for the evolutionary process, and
adaptive trait for the bodily part or function (the product), one may
distinguish the two different senses of the word.
Adaptation is one of the two main processes that explain the observed
diversity of species, such as the different species of Darwin's
finches. The other process is speciation, in which new species arise,
typically through reproductive isolation. A favourite example
used today to study the interplay of adaptation and speciation is the
evolution of cichlid fish in African lakes, where the question of
reproductive isolation is complex.
Adaptation is not always a simple matter where the ideal phenotype
evolves for a given external environment. An organism must be viable
at all stages of its development and at all stages of its evolution.
This places constraints on the evolution of development, behaviour,
and structure of organisms. The main constraint, over which there has
been much debate, is the requirement that each genetic and phenotypic
change during evolution should be relatively small, because
developmental systems are so complex and interlinked. However, it is
not clear what "relatively small" should mean, for example polyploidy
in plants is a reasonably common large genetic change. The origin
of eukaryotic endosymbiosis is a more dramatic example.
All adaptations help organisms survive in their ecological niches.
The adaptive traits may be structural, behavioural or physiological.
Structural adaptations are physical features of an organism, such as
shape, body covering, armament, and internal organization. Behavioural
adaptations are inherited systems of behaviour, whether inherited in
detail as instincts, or as a neuropsychological capacity for learning.
Examples include searching for food, mating, and vocalizations.
Physiological adaptations permit the organism to perform special
functions such as making venom, secreting slime, and phototropism),
but also involve more general functions such as growth and
development, temperature regulation, ionic balance and other aspects
Adaptation affects all aspects of the life of an
The following definitions are given by the evolutionary biologist
Adaptation is the evolutionary process whereby an organism becomes
better able to live in its habitat or habitats.
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.
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.
What adaptation is not
Some generalists, such as birds, have the flexibility to adapt to
Adaptation differs from flexibility, acclimatization, and learning.
Flexibility deals with the relative capacity of an organism to
maintain itself in different habitats: its degree of specialization.
Acclimatization describes automatic physiological adjustments during
life; learning means improvement in behavioral performance during
life. These terms are preferred to adaptation for changes during
life which are not inherited by the next generation.
Flexibility stems from phenotypic plasticity, the ability of an
organism with a given genotype to change its phenotype in response to
changes in its habitat, or to move to a different habitat. The
degree of flexibility is inherited, and varies between individuals. A
highly specialized animal or plant lives only in a well-defined
habitat, eats a specific type of food, and cannot survive if its needs
are not met. Many herbivores are like this; extreme examples are
koalas which depend on Eucalyptus, and giant pandas which require
bamboo. A generalist, on the other hand, eats a range of food, and can
survive in many different conditions. Examples are humans, rats, crabs
and many carnivores. The tendency to behave in a specialized or
exploratory manner is inherited—it is an adaptation. Rather
different is developmental flexibility: "An animal or plant is
developmentally flexible if when it is raised in or transferred to new
conditions, it changes in structure so that it is better fitted to
survive in the new environment," writes evolutionary biologist John
If humans move to a higher altitude, respiration and physical exertion
become a problem, but after spending time in high altitude conditions
they acclimatize to the reduced partial pressure of oxygen, such as by
producing more red blood cells. The ability to acclimatize is an
adaptation, but the acclimatization itself is not.
down, but deaths from some tropical diseases also go down. Over a
longer period of time, some people are better able to reproduce at
high altitudes than others. They contribute more heavily to later
generations, and gradually by natural selection the whole population
becomes adapted to the new conditions. This has demonstrably occurred,
as the observed performance of long-term communities at higher
altitude is significantly better than the performance of new arrivals,
even when the new arrivals have had time to acclimatize.
Adaptedness and fitness
In this sketch of a fitness landscape, a population can evolve by
following the arrows to the adaptive peak at point B, and the points A
and C are local optima where a population could become trapped.
Fitness (biology) and Fitness landscape
There is a relationship between adaptedness and the concept of fitness
used in population genetics. Differences in fitness between genotypes
predict the rate of evolution by natural selection. Natural selection
changes the relative frequencies of alternative phenotypes, insofar as
they are heritable. However, a phenotype with high adaptedness may
not have high fitness. Dobzhansky mentioned the example of the
Californian redwood, which is highly adapted, but a relict species in
danger of extinction.
Elliott Sober commented that adaptation was
a retrospective concept since it implied something about the history
of a trait, whereas fitness predicts a trait's future.
1. Relative fitness. The average contribution to the next generation
by a genotype or a class of genotypes, relative to the contributions
of other genotypes in the population. This is also known as
Darwinian fitness, selection coefficient, and other terms.
2. Absolute fitness. The absolute contribution to the next generation
by a genotype or a class of genotypes. Also known as the Malthusian
parameter when applied to the population as a whole.
3. Adaptedness. The extent to which a phenotype fits its local
ecological niche. Researchers can sometimes test this through a
Sewall Wright proposed that populations occupy adaptive peaks on a
fitness landscape. To evolve to another, higher peak, a population
would first have to pass through a valley of maladaptive intermediate
stages, and might be "trapped" on a peak that is not optimally
A large diversity of genome DNAs in a species is the basis for
adaptation and differentiation. A large population is needed to carry
sufficient diversity. According to the misrepair-accumulation aging
theory, The misrepair mechanism is important in maintaining a
sufficient number of individuals in a species. misrepair is a way
of repair for increasing the surviving chance of an organism when it
has severe injuries. Without misrepairs, no individual could survive
to reproduction age. Thus misrepair mechanism is an essential
mechanism for the survival of a species and for maintaining the number
of individuals. Although individuals die from aging, genome DNAs are
being recopied and transmitted by individuals generation by
generation. In addition, the
DNA misrepairs in germ cells contribute
also to the diversity of genome DNAs.
Adaptation is the heart and soul of evolution.
— Niles Eldredge, Reinventing Darwin: The Great Debate at the High
Table of Evolutionary Theory
Changes in habitat
Before Darwin, adaptation was seen as a fixed relationship between an
organism and its habitat. It was not appreciated that as the climate
changed, so did the habitat; and as the habitat changed, so did the
biota. Also, habitats are subject to changes in their biota: for
example, invasions of species from other areas. The relative numbers
of species in a given habitat are always changing. Change is the rule,
though much depends on the speed and degree of the change. When the
habitat changes, three main things may happen to a resident
population: habitat tracking, genetic change or extinction. In fact,
all three things may occur in sequence. Of these three effects only
genetic change brings about adaptation. When a habitat changes, the
resident population typically moves to more suitable places; this is
the typical response of flying insects or oceanic organisms, which
have wide (though not unlimited) opportunity for movement. This
common response is called habitat tracking. It is one explanation put
forward for the periods of apparent stasis in the fossil record (the
punctuated equilibrium theory).
Genetic change occurs in a population when natural selection and
mutations act on its genetic variability. The first pathways of
enzyme-based metabolism may have been parts of purine nucleotide
metabolism, with previous metabolic pathways being part of the ancient
RNA world. By this means, the population adapts genetically to its
circumstances. Genetic changes may result in visible structures,
or may adjust physiological activity in a way that suits the habitat.
Habitats and biota do frequently change. Therefore, it follows that
the process of adaptation is never finally complete. Over time, it
may happen that the environment changes little, and the species comes
to fit its surroundings better and better. On the other hand, it may
happen that changes in the environment occur relatively rapidly, and
then the species becomes less and less well adapted. Seen like this,
adaptation is a genetic tracking process, which goes on all the time
to some extent, but especially when the population cannot or does not
move to another, less hostile area. Given enough genetic change, as
well as specific demographic conditions, an adaptation may be enough
to bring a population back from the brink of extinction in a process
called evolutionary rescue. It should be noted that adaptation does
affect, to some extent, every species in a particular
Leigh Van Valen
Leigh Van Valen thought that even in a stable environment, competing
species constantly had to adapt to maintain their relative standing.
This became known as the Red Queen hypothesis, as seen in
Main article: Co-adaptation
Pollinating insects are co-adapted with flowering plants.
In coevolution, where the existence of one species is tightly bound up
with the life of another species, new or 'improved' adaptations which
occur in one species are often followed by the appearance and spread
of corresponding features in the other species. These co-adaptational
relationships are intrinsically dynamic, and may continue on a
trajectory for millions of years, as has occurred in the relationship
between flowering plants and pollinating insects.
Main article: Mimicry
A and B show real wasps; the rest are Batesian mimics: three
hoverflies and one beetle.
Bates' work on Amazonian butterflies led him to develop the first
scientific account of mimicry, especially the kind of mimicry which
bears his name: Batesian mimicry. This is the mimicry by a
palatable species of an unpalatable or noxious species, gaining a
selective advantage. A common example seen in temperate gardens is the
hoverfly, many of which—though bearing no sting—mimic the warning
coloration of hymenoptera (wasps and bees). Such mimicry does not need
to be perfect to improve the survival of the palatable species.
Bates, Wallace and
Fritz Müller believed that Batesian and Müllerian
mimicry provided evidence for the action of natural selection, a view
which is now standard amongst biologists.
It is a profound truth that Nature does not know best; that genetical
evolution... is a story of waste, makeshift, compromise and blunder.
— Peter Medawar, The Future of Man
All adaptations have a downside: horse legs are great for running on
grass, but they can't scratch their backs; mammals' hair helps
temperature, but offers a niche for ectoparasites; the only flying
penguins do is under water. Adaptations serving different functions
may be mutually destructive. Compromise and makeshift occur widely,
not perfection. Selection pressures pull in different directions, and
the adaptation that results is some kind of compromise.
Since the phenotype as a whole is the target of selection, it is
impossible to improve simultaneously all aspects of the phenotype to
the same degree.
— Ernst Mayr, The Growth of Biological Thought: Diversity,
Evolution, and Inheritance
Consider the antlers of the Irish elk, (often supposed to be far too
large; in deer antler size has an allometric relationship to body
size). Obviously, antlers serve positively for defence against
predators, and to score victories in the annual rut. But they are
costly in terms of resource. Their size during the last glacial period
presumably depended on the relative gain and loss of reproductive
capacity in the population of elks during that time. As another
example, camouflage to avoid detection is destroyed when vivid
coloration is displayed at mating time. Here the risk to life is
counterbalanced by the necessity for reproduction.
Stream-dwelling salamanders, such as
Caucasian salamander or
Gold-striped salamander have very slender, long bodies, perfectly
adapted to life at the banks of fast small rivers and mountain brooks.
Elongated body protects their larvae from being washed out by current.
However, elongated body increases risk of desiccation and decreases
dispersal ability of the salamanders; it also negatively affects their
fecundity. As a result, fire salamander, less perfectly adapted to the
mountain brook habitats, is in general more successful, have a higher
fecundity and broader geographic range.
An Indian peacock's train
in full display
The peacock's ornamental train (grown anew in time for each mating
season) is a famous adaptation. It must reduce his maneuverability and
flight, and is hugely conspicuous; also, its growth costs food
resources. Darwin's explanation of its advantage was in terms of
sexual selection: "This depends on the advantage which certain
individuals have over other individuals of the same sex and species,
in exclusive relation to reproduction." The kind of sexual
selection represented by the peacock is called 'mate choice,' with an
implication that the process selects the more fit over the less fit,
and so has survival value. The recognition of sexual selection was
for a long time in abeyance, but has been rehabilitated.
The conflict between the size of the human foetal brain at birth,
(which cannot be larger than about 400 cm3, else it will not get
through the mother's pelvis) and the size needed for an adult brain
(about 1400 cm3), means the brain of a newborn child is quite
immature. The most vital things in human life (locomotion, speech)
just have to wait while the brain grows and matures. That is the
result of the birth compromise. Much of the problem comes from our
upright bipedal stance, without which our pelvis could be shaped more
suitably for birth. Neanderthals had a similar problem.
As another example, the long neck of a giraffe is a burden and a
blessing. The neck of a giraffe can be up to 2 m (6 ft
7 in) in length. This neck can be used for inter-species
competition or for foraging on tall trees where shorter herbivores
cannot reach. However, as previously stated, there is always a
trade-off. This long neck is heavy and it adds to the body mass of a
giraffe, so the giraffe needs an abundance of nutrition to provide for
this costly adaptation.
Shifts in function
Adaptation and function are two aspects of one problem.
— Julian Huxley, Evolution: The Modern Synthesis
Pre-adaptation occurs when a population has characteristics which by
chance are suited for a set of conditions not previously experienced.
For example, the polyploid cordgrass
Spartina townsendii is better
adapted than either of its parent species to their own habitat of
saline marsh and mud-flats. Among domestic animals, the White
Leghorn chicken is markedly more resistant to vitamin B1 deficiency
than other breeds; on a plentiful diet this makes no difference, but
on a restricted diet this preadaptation could be decisive.
Pre-adaptation may arise because a natural population carries a huge
quantity of genetic variability. In diploid eukaryotes, this is a
consequence of the system of sexual reproduction, where mutant alleles
get partially shielded, for example, by genetic dominance.
Microorganisms, with their huge populations, also carry a great deal
of genetic variability. The first experimental evidence of the
pre-adaptive nature of genetic variants in microorganisms was provided
Salvador Luria and
Max Delbrück who developed the Fluctuation
Test, a method to show the random fluctuation of pre-existing genetic
changes that conferred resistance to bacteriophages in Escherichia
Co-option of existing traits: exaptation
Sinosauropteryx fossil, a dinosaur with feathers, used for insulation,
Main article: Exaptation
Features that now appear as adaptations sometimes arose by co-option
of existing traits, evolved for some other purpose. The classic
example is the ear ossicles of mammals, which we know from
paleontological and embryological evidence originated in the upper and
lower jaws and the hyoid bone of their synapsid ancestors, and further
back still were part of the gill arches of early fish. The
word exaptation was coined to cover these common evolutionary shifts
in function. The flight feathers of birds evolved from the much
earlier feathers of dinosaurs, which might have been used for
insulation or for display.
Spandrel (biology) and Vestigiality
Some traits do not appear to be adaptive, that is, they have a neutral
or deleterious effect on fitness in the current environment. Because
genes have pleiotropic effects, not all traits may be functional: they
may be what
Stephen Jay Gould
Stephen Jay Gould and
Richard Lewontin called spandrels,
features brought about by neighbouring adaptations, like the
triangular areas under neighbouring arches in architecture which began
as functionless features.
Another possibility is that a trait may have been adaptive at some
point in an organism's evolutionary history, but a change in habitats
caused what used to be an adaptation to become unnecessary or even
maladapted. Such adaptations are termed vestigial. Many organisms have
vestigial organs, which are the remnants of fully functional
structures in their ancestors. As a result of changes in lifestyle the
organs became redundant, and are either not functional or reduced in
functionality. Since any structure represents some kind of cost to the
general economy of the body, an advantage may accrue from their
elimination once they are not functional. Examples: wisdom teeth in
humans; the loss of pigment and functional eyes in cave fauna; the
loss of structure in endoparasites.
Extinction and coextinction
Extinction and Coextinction
If a population cannot move or change sufficiently to preserve its
long-term viability, then obviously, it will become extinct, at least
in that locale. The species may or may not survive in other locales.
Species extinction occurs when the death rate over the entire species
exceeds the birth rate for a long enough period for the species to
disappear. It was an observation of Van Valen that groups of species
tend to have a characteristic and fairly regular rate of
Just as there is co-adaptation, there is also coextinction, the loss
of a species due to the extinction of another with which it is
coadapted, as with the extinction of a parasitic insect following the
loss of its host, or when a flowering plant loses its pollinator, or
when a food chain is disrupted.
Teleology in biology
Adaptation raises philosophical issues concerning how biologists speak
of function and purpose, as this carries implications of evolutionary
history – that a feature evolved by natural selection for a specific
reason – and potentially of supernatural intervention – that
features and organisms exist because of a deity's conscious
intentions. In his biology,
Aristotle introduced teleology to
describe the adaptedness of organisms, but without accepting the
supernatural intention built into Plato's thinking, which Aristotle
rejected. Modern biologists continue to face the same
difficulty. On the one hand, adaptation is
obviously purposeful: natural selection chooses what works and
eliminates what does not. On the other hand, biologists want to deny
conscious purpose in evolution. The dilemma gave rise to a famous joke
by the evolutionary biologist Haldane: "
Teleology is like a mistress
to a biologist: he cannot live without her but he's unwilling to be
seen with her in public.'"
David Hull commented that Haldane's
mistress "has become a lawfully wedded wife. Biologists no longer feel
obligated to apologize for their use of teleological language; they
Adaptive evolution in the human genome
Neutral theory of molecular evolution
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Evolutionary developmental biology
Hierarchy of life
Ecosystem > Community (Biocoenosis)
Organism > Organ system
> Organ > Tissue > Cell > Organelle
Biomolecular complex >
Biomolecule) > Atom
Earliest known life forms
Plant morphology terms
Evolutionary history of life
Index of evolutionary biology articles
Outline of evolution
Timeline of evolution
Earliest known life forms
Evidence of common descent
Last universal common ancestor
Origin of life
Evolutionary developmental biology
dolphins and whales
Programmed cell death
Life cycles/nuclear phases
Tempo and modes
Renaissance and Enlightenment
Transmutation of species
On the Origin of Species
History of paleontology
The eclipse of Darwinism
History of molecular evolution
Extended evolutionary synthesis
Teleology in biology
Species problem ·
Evidence of common descent
Allopatric (Peripatric ·
Founder effect · Centrifugal)
Parapatric (Clines · Ring species)
Hybrid speciation (
Polyploidy · Recombination)
Speciation in taxa
Punctuated equilibrium · Macroevolution