Adaptation

In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural
selection 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 trait or adaptive trait, with a
functional role in each individual organism, that is maintained and has evolved through natural selection.

Historically, adaptation has been described from the time of the ancient Greek philosophers such as
Empedocles and Aristotle. In 18th and 19th century natural theology, adaptation was taken as evidence for
the existence of a deity. Charles Darwin proposed instead that it was explained by natural selection.

Adaptation is related to biological fitness, which governs the rate of evolution as measured by change in
allele frequencies. Often, two or more species co-adapt and co-evolve as they develop adaptations that
interlock with those of the other species, such as with flowering plants and pollinating insects. In mimicry,
species evolve to resemble other species; in Müllerian mimicry this is a mutually beneficial co-evolution as
each of a group of strongly defended species (such as wasps able to sting) come to advertise their defenses
in the same way. Features evolved for one purpose may be co-opted for a different one, as when the
insulating feathers of dinosaurs were co-opted for bird flight.

Adaptation is a major topic in the philosophy of biology, as it concerns function and purpose (teleology).
Some biologists try to avoid terms which imply purpose in adaptation, not least because it suggests a deity's
intentions, but others note that adaptation is necessarily purposeful.

History
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 (a purpose), but thought that it "came about naturally, since such things
survived." Aristotle did believe in final causes, but assumed that species were fixed.[1]

In natural theology, adaptation was interpreted as the work of a deity and as evidence for the existence of
God.[2] 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 satire Dr. Pangloss[3] is a parody of this optimistic idea, and David Hume also argued
against design.[4] Charles Darwin broke with the tradition by emphasising the flaws and limitations which
occurred in the animal and plant worlds.[5]

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.[6] 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.[7]

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
and Theodosius Dobzhansky produced
evidence that natural selection was not only
the 'engine' behind adaptation, but was a
much stronger force than had previously
been thought.[8][9][10]

General principles

The significance of an
adaptation can only be
understood in relation to the
total biology of the species.
The second of Jean-Baptiste Lamarck's two factors (the first
— Julian Huxley, being a complexifying force) was an adaptive force that
causes animals with a given body plan to adapt to
Evolution: The Modern
circumstances by inheritance of acquired characteristics,
Synthesis[11] creating a diversity of species and genera.

What adaptation is

Adaptation is primarily a process rather than a physical form or part of a body.[12] 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.[13] 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.[14][15][16][17]

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.[18][19] An example widely 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.[20][21]

Adaptation is not always a simple matter where the ideal phenotype evolves for a given 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.[22] The origin of eukaryotic endosymbiosis is a more dramatic
example.[23]

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 of homeostasis. Adaptation affects all aspects of the life of an organism.[24]

The following definitions are given by the evolutionary biologist Theodosius Dobzhansky:

1. Adaptation is the evolutionary process whereby an organism becomes better able to

live in its habitat or habitats.[25][26][27]
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.[28]
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.[29]

What adaptation is not

Adaptation differs from flexibility, acclimatization, and learning, all

of which are changes during life which are not inherited. 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;[30]
learning means improvement in behavioural performance during
life.[31]

Flexibility stems from phenotypic plasticity, the ability of an Some generalists, such as birds,
organism with a given genotype (genetic type) to change its
have the flexibility to adapt to urban
phenotype (observable characteristics) in response to changes in its areas.
habitat, or to move to a different habitat.[32][33] 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 the evolutionary biologist John Maynard Smith.[34]

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. The reproductive rate declines, 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.[35]

Adaptedness and fitness

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.[36] 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.[25] 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.[37]

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.[38]
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.[36][39]
3. Adaptedness. The extent to which a phenotype fits its local ecological niche.
Researchers can sometimes test this through a reciprocal transplant.[40]

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 adapted.[41]

Types In this sketch of a fitness

landscape, a population can evolve
by following the arrows to the
adaptive peak at point B, and the
Adaptation is the heart and soul of evolution.
points A and C are local optima
where a population could become
— Niles Eldredge, Reinventing Darwin: The Great
trapped.
Debate at the High Table of Evolutionary
Theory[42]

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.[43] 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).[44]

Genetic change
Without mutation, the ultimate source of all genetic variation, there would be no genetic changes and no
subsequent adaptation through evolution by natural selection. Genetic change occurs in a population when
mutation increases or decreases in its initial frequency followed by random genetic drift, migration,
recombination or natural selection act on this genetic variation.[45] One example is that the first pathways of
enzyme-based metabolism at the very origin of life on Earth may have been co-opted components of the
already-existing purine nucleotide metabolism, a metabolic pathway that evolved in an ancient RNA world.
The co-option requires new mutations and through natural selection, the population then adapts genetically
to its present circumstances.[10] Genetic changes may result in entirely new or gradual change to visible
structures, or they may adjust physiological activity in a way that suits the habitat. The varying shapes of
the beaks of Darwin's finches, for example, are driven by adaptive mutations in the ALX1 gene.[46] The
coat color of different wild mouse species matches their environments, whether black lava or light sand,
owing to adaptive mutations in the melanocortin 1 receptor and other melanin pathway genes.[47][48]
Physiological resistance to the heart poisons (cardiac glycosides) that monarch butterflies store in their
bodies to protect themselves from predators[49][50] are driven by adaptive mutations in the target of the
poison, the sodium pump, resulting in target site insensitivity.[51][52][53] These same adaptive mutations and
similar changes at the same amino acid sites were found to evolve in a parallel manner in distantly related
insects that feed on the same plants, and even in a bird that feeds on monarchs through convergent
evolution, a hallmark of adaptation.[54][55] Convergence at the gene-level across distantly related species
can arise because of evolutionary constraint.[56]

Habitats and biota do frequently change over time and space. Therefore, it follows that the process of
adaptation is never fully complete.[57] Over time, it may happen that the environment changes little, and the
species comes to fit its surroundings better and better, resulting in stabilizing selection. On the other hand, it
may happen that changes in the environment occur suddenly, and then the species becomes less and less
well adapted. The only way for it to climb back up that fitness peak is via the introduction of new genetic
variation for natural selection to act upon. 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. Adaptation does affect, to some extent, every species in a particular ecosystem.[58][59]

Leigh Van Valen thought that even in a stable environment, because of antagonistic species interactions and
limited resources, a species must constantly had to adapt to maintain its relative standing. This became
known as the Red Queen hypothesis, as seen in host-parasite interactions.[60]

Existing genetic variation and mutation were the traditional sources of material on which natural selection
could act. In addition, horizontal gene transfer is possible between organisms in different species, using
mechanisms as varied as gene cassettes, plasmids, transposons and viruses such as
bacteriophages.[61][62][63]

Co-adaptation

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. In other words, each species triggers reciprocal natural selection
in the other. 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.[64][65]

Mimicry

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.[66] This is the mimicry by a Pollinating insects are co-adapted
palatable species of an unpalatable or noxious species (the model), with flowering plants.
gaining a selective advantage as predators avoid the model and
therefore also the mimic. Mimicry is thus an anti-predator
adaptation. A common example seen in temperate gardens is the
hoverfly (Syrphidae), many of which—though bearing no sting—
mimic the warning coloration of aculeate Hymenoptera (wasps and
bees). Such mimicry does not need to be perfect to improve the
survival of the palatable species.[67]

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.[68][69][70]

Trade-offs

It is a profound truth that Nature does not know best;

Images A and B show real wasps;
that genetical evolution... is a story of waste, makeshift,
the others show Batesian mimics:
compromise and blunder.
three hoverflies and one beetle.

— Peter Medawar, The Future of Man[71]

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.[72]

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[73]

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 resources. 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.[74] 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.[75]

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.[76]

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."[77] The kind
of sexual selection represented by the peacock is called 'mate
choice,' with an implication that the process selects the more fit over
An Indian peacock's train the less fit, and so has survival value.[78] The recognition of sexual
in full display selection was for a long time in abeyance, but has been
rehabilitated.[79]

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.[80][81][82]

As another example, the long neck of a giraffe brings benefits but at a cost. The neck of a giraffe can be up
to 2 m (6 ft 7 in) in length.[83] The benefits are that it can be used for inter-species competition or for
foraging on tall trees where shorter herbivores cannot reach. The cost is that a long neck is heavy and adds
to the animal's body mass, requiring additional energy to build the neck and to carry its weight around.[84]

Shifts in function

Adaptation and function are two aspects of one problem.

— Julian Huxley, Evolution: The Modern Synthesis[85]

Pre-adaptation

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.[86] 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.[87]

Pre-adaptation may arise because a natural population carries a huge quantity of genetic variability.[88] In
diploid eukaryotes, this is a consequence of the system of sexual reproduction, where mutant alleles get
partially shielded, for example, by genetic dominance.[89] 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 by 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 coli.[90] The word is controversial because it is teleological and
the entire concept of natural selection depends on the presence of genetic variation, regardless of the
population size of a species in question.

Co-option of existing traits: 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.[91][92] The word exaptation was
coined to cover these common evolutionary shifts in
function.[93] The flight feathers of birds evolved from the The feathers of Sinosauropteryx, a dinosaur
much earlier feathers of dinosaurs,[94] which might have with feathers, were used for insulation or
display, making them an exaptation for
been used for insulation or for display.[95][96]
flight.

Niche construction
Animals including earthworms, beavers and humans use some of their adaptations to modify their
surroundings, so as to maximize their chances of surviving and reproducing. Beavers create dams and
lodges, changing the ecosystems of the valleys around them. Earthworms, as Darwin noted, improve the
topsoil in which they live by incorporating organic matter. Humans have constructed extensive civilizations
with cities in environments as varied as the Arctic and hot deserts. In all three cases, the construction and
maintenance of ecological niches helps drive the continued selection of the genes of these animals, in an
environment that the animals have modified.[97]

Non-adaptive traits
Some traits do not appear to be adaptive as they have a neutral or deleterious effect on fitness in the current
environment. Because genes often have pleiotropic effects, not all traits may be functional: they may be
what Stephen Jay Gould and Richard Lewontin called spandrels, features brought about by neighbouring
adaptations, on the analogy with the often highly decorated triangular areas between pairs of arches in
architecture, which began as functionless features.[98]

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.[99]

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 extinction.[100]

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.[101][102]

Origin of adaptive capacities

The first stage in the evolution of life on earth is often hypothesized to be the RNA world in which short
self-replicating RNA molecules proliferated before the evolution of DNA and proteins. By this hypothesis,
life started when RNA chains began to self-replicate, initiating the three mechanisms of Darwinian
selection: heritability, variation of type, and competition for resources. The fitness of an RNA replicator (its
per capita rate of increase) would likely have been a function of its intrinsic adaptive capacities, determined
by its nucleotide sequence, and the availability of resources.[103][104] The three primary adaptive capacities
may have been: (1) replication with moderate fidelity, giving rise to heritability while allowing variation of
type, (2) resistance to decay, and (3) acquisition of resources.[103][104] These adaptive capacities would
have been determined by the folded configurations of the RNA replicators resulting from their nucleotide
sequences.

Philosophical issues
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.[107][108] 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.[109][110] Modern biologists continue to face the same difficulty.[111][112][113][114][115] On the one
hand, adaptation is obviously purposeful: natural selection chooses what works and eliminates what does
not. On the other hand, biologists by and large reject 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 flaunt it."[116] Ernst
Mayr stated that "adaptedness... is a posteriori result rather than an
a priori goal-seeking", meaning that the question of whether
something is an adaptation can only be determined after the
event.[117]

See also
Adaptive evolution in the human genome
Adaptive memory
Adaptive mutation
Adaptive system
Anti-predator adaptation
"Behaviour with a purpose": a young
Body reactivity springbok stotting. A biologist might
Ecological trap argue that this has the function of
Evolutionary pressure signalling to predators, helping the
Evolvability springbok to survive and allowing it
Intragenomic conflict to reproduce.[105][106]

Neutral theory of molecular evolution

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