Adaptation

This article is about the evolutionary process. For other

uses, see Adaptation (disambiguation).
Not to be confused with Adoption or Acclimatization.
In biology, an adaptation, also called an adaptive trait,
is a trait with a current functional role in the life history
of an organism that is maintained and evolved by means
of natural selection. Adaptation refers to both the cur-
rent state of being adapted and to the dynamic evolu-
tionary process that leads to the adaptation. Adaptations
contribute to the tness and survival of individuals. Or-
ganisms face a succession of environmental challenges as
they grow and develop and are equipped with an adaptive
plasticity as the phenotype of traits develop in response
to the imposed conditions. The developmental norm of
reaction for any given trait is essential to the correction
of adaptation as it aords a kind of biological insurance
or resilience to varying environments.
1 General principles
The signicance of an adaptation can only
be understood in relation to the total biology of
the species.
Julian Huxley
[1]
Adaptation is, rst of all, a process, rather than a physical
part of a body.
[2]
An internal parasite (such as a uke)
can illustrate the distinction: such a parasite may have a
very simple bodily structure, but nevertheless the organ-
ism is highly adapted to its specic environment. From
this we see that adaptation is not just a matter of visi-
ble traits: in such parasites critical adaptations take place
in the life-cycle, which is often quite complex.
[3]
How-
ever, as a practical term, adaptation is often used for the
product: those features of a species which result from the
process. Many aspects of an animal or plant can be cor-
rectly called adaptations, though there are always some
features whose function is in doubt. By using the term
adaptation for the evolutionary process, and adaptive
trait for the bodily part or function (the product), the
two senses of the word may be distinguished.
[4][5][6][7]
Adaptation is one of the two main processes that ex-
plain the diverse species we see in biology, such as
the dierent species of Darwins nches. The other is
speciation (species-splitting or cladogenesis), caused by
geographical isolation or some other mechanism.
[8][9]
A
favorite example used today to study the interplay of
adaptation and speciation is the evolution of cichlid sh
in African lakes, where the question of reproductive iso-
lation is much more complex.
[10][11]
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 struc-
ture of organisms. The main constraint, over which there
has been much debate, is the requirement that each ge-
netic 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.
[12]
The origin of eukaryotic symbiosis is a more dramatic
example.
[13]
All adaptations help organisms survive in their ecological
niches.
[14]
These adaptive traits may be structural, behav-
ioral or physiological. Structural adaptations are phys-
ical features of an organism (shape, body covering, ar-
mament; and also the internal organization). Behavioural
adaptations are composed of inherited behaviour chains
and/or the ability to learn: behaviours may be inherited
in detail (instincts), or a capacity for learning may be
inherited (see neuropsychology). Examples: searching
for food, mating, vocalizations. Physiological adaptations
permit the organism to perform special functions (for in-
stance, making venom, secreting slime, phototropism);
but also more general functions such as growth and
development, temperature regulation, ionic balance and
other aspects of homeostasis. Adaptation, then, aects
1
2 2 BRIEF HISTORY
all aspects of the life of an organism.
1.1 Denitions
The following denitions are mainly due to Theodosius
Dobzhansky.
1. Adaptation is the evolutionary process
whereby an organism becomes better able to
live in its habitat or habitats.
[15]
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.
[16]
3. An adaptive trait is an aspect of the develop-
mental pattern of the organism which enables
or enhances the probability of that organism
surviving and reproducing.
[17]
1.2 Adaptedness and tness
Main article: Fitness (biology)
From the above denitions, it is clear that there is
a relationship between adaptedness and tness (a key
population genetics concept). Dierences in tness be-
tween genotypes predict the rate of evolution by natu-
ral selection. Natural selection changes the relative fre-
quencies of alternative phenotypes, insofar as they are
heritable.
[18]
Although the two are connected, the one
does not imply the other: a phenotype with high adapted-
ness may not have high tness. Dobzhansky mentioned
the example of the Californian redwood, which is highly
adapted, but a relict species in danger of extinction.
[15]
Elliott Sober commented that adaptation was a retrospec-
tive concept since it implied something about the history
of a trait, whereas tness predicts a traits future.
[19]p210
1. Relative tness. 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.
[20]p552
This
is also known as Darwinian tness, selective co-
ecient, and other terms.
2. Absolute tness. 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.
[18]
3. Adaptedness. The extent to which a phe-
notype ts its local ecological niche. This can
sometimes be tested through a reciprocal trans-
plant experiment.
2 Brief history
Main article: History of evolutionary thought
Adaptation is a fact of life that has been accepted by many
of the great thinkers who have tackled the world of liv-
ing organisms. It is their explanations of how adaptation
arises that separates these thinkers. A few of the most
signicant ideas:
[21]
Empedocles did not believe that adaptation required
a nal cause (~ purpose), but came about naturally,
since such things survived. Aristotle, however, did
believe in nal causes.
In natural theology, adaptation was interpreted as
the work of a deity, even as evidence for the exis-
tence of God.
[22]
William Paley believed that organ-
isms were perfectly adapted to the lives they lead,
an argument that shadowed Leibniz, who had ar-
gued that God had brought about the best of all
possible worlds. Voltaire's Dr Pangloss
[23]
is a par-
ody of this optimistic idea, and Hume also argued
against design.
[24]
The Bridgewater Treatises are a
product of natural theology, though some of the au-
thors managed to present their work in a fairly neu-
tral manner. The series was lampooned by Robert
Knox, who held quasi-evolutionary views, as the Bil-
gewater Treatises. Darwin broke with the tradition
by emphasising the aws and limitations which oc-
curred in the animal and plant worlds.
[25]
Lamarck's is a proto-evolutionary theory of the
inheritance of acquired traits, whose main purpose
is to explain adaptations by natural means.
[26]
He
proposed a tendency for organisms to become more
complex, moving up a ladder of progress, plus the
inuence of circumstances, usually expressed as
use and disuse. His evolutionary ideas, and those
of Georoy, fail because they cannot be reconciled
3.1 Changes in habitat 3
Lamarck
with heredity. This was known even before Mendel
by medical men interested in human races (Wells,
Lawrence), and especially by Weismann.
Many other students of natural history, such as Buon,
accepted adaptation, and some also accepted evolution,
without voicing their opinions as to the mechanism. This
illustrates the real merit of Darwin and Wallace, and
secondary gures such as Bates, for putting forward a
mechanism whose signicance had only been glimpsed
previously. A century later, experimental eld studies
and breeding experiments by people such as Ford and
Dobzhansky produced evidence that natural selection was
not only the 'engine' behind adaptation, but was a much
stronger force than had previously been thought.
[27][28][29]
3 Types of adaptations
Adaptation is the heart and soul of evolu-
tion. -Niles Eldredge
[30]
3.1 Changes in habitat
Before Darwin, adaptation was seen as a xed relation-
ship between an organism and its habitat. It was not ap-
preciated that as the climate changed, so did the habitat;
and as the habitat changed, so did the biota. Also, habi-
tats are subject to changes in their biota: for example,
invasions of species from other areas. The relative num-
bers 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 se-
quence. Of these three eects, only genetic change brings
about adaptation.
3.1.1 Habitat tracking
When a habitat changes, the most common thing to hap-
pen is that the resident population moves to another lo-
cale which suits it; this is the typical response of ying
insects or oceanic organisms, who have wide (though not
unlimited) opportunity for movement.
[31]
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 thesis).
[32]
3.1.2 Genetic change
Genetic change is what occurs in a population when
natural selection acts on the genetic variability of the
population; moreover, some mutations may create ge-
netic variation that will lead to diering characteris-
tics of ospring and hence abet adaptation.
[33]
The rst
pathways of enzyme-based metabolism may have been
parts of purine nucleotide metabolism, with previous
metabolic pathways being part of the ancient RNAworld.
By this means, the population adapts genetically to its
circumstances.
[34]
Genetic changes may result in visible
structures, or may adjust physiological activity in a way
that suits the habitat.
It is now clear that habitats and biota do frequently
change. Therefore, it follows that the process of adapta-
tion is never nally complete.
[35]
Over time, it may hap-
pen that the environment changes little, and the species
comes to t its surroundings better and better. On the
4 3 TYPES OF ADAPTATIONS
other hand, it may happen that changes in the environ-
ment occur relatively rapidly, and then the species be-
comes less and less well adapted. Seen like this, adap-
tation 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.
Also, to a greater or lesser extent, the process aects ev-
ery species in a particular ecosystem.
[36][37]
Van Valen thought that even in a stable environment,
competing species had to constantly adapt to maintain
their relative standing. This became known as the Red
Queens hypothesis.
3.2 Intimate relationships: co-adaptations
Main article: Co-adaptation
In co-evolution, where the existence of one species is
tightly over 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 corre-
sponding features in the other species. There are many
examples of this; the idea emphasises that the life and
death of living things is intimately connected, not just
with the physical environment, but with the life of other
species. These relationships are intrinsically dynamic,
and may continue on a trajectory for millions of years,
as has the relationship between owering plants and in-
sects (pollination).
Pollinator constancy: these honeybees selectively visit
owers from only one species, as can be seen by the
colour of the pollen in their baskets:

Co-extinction
Infection-resistance
Mimicry
Mutualism
Parasite-host
Pollination syndrome
Predator-prey
Symbiosis
The gut contents, wing structures, and mouthpart mor-
phologies of fossilized beetles and ies suggest that they
acted as early pollinators. The association between
beetles and angiosperms during the early Cretaceous pe-
riod led to parallel radiations of angiosperms and in-
sects into the late Cretaceous. The evolution of nectaries
in late Cretaceous owers signals the beginning of the
mutualism between hymenopterans and angiosperms.
[38]
3.2.1 Mimicry
Main article: Mimicry
Henry Walter Bates' work on Amazonian butteries led
A and B show real wasps; the rest are mimics: three hoveries
and one beetle.
him to develop the rst scientic account of mimicry,
especially the kind of mimicry which bears his name:
Batesian mimicry.
[39]
This is the mimicry by a palatable
species of an unpalatable or noxious species. A common
example seen in temperate gardens is the hover-y, many
3.4 Compromise and conict between adaptations 5
of which though bearing no sting mimic the warn-
ing colouration of hymenoptera (wasps and bees). Such
mimicry does not need to be perfect to improve the sur-
vival of the palatable species.
[40]
Bates, Wallace and Mller believed that Batesian and
Mllerian mimicry provided evidence for the action of
natural selection, a view which is now standard amongst
biologists.
[41]
All aspects of this situation can be, and have
been, the subject of research.
[42]
Field and experimental
work on these ideas continues to this day; the topic con-
nects strongly to speciation, genetics and development.
[43]
3.3 The basic machinery: internal adapta-
tions
There are some important adaptations to do with the over-
all coordination of the systems in the body. Such adap-
tations may have signicant consequences. Examples,
in vertebrates, would be temperature regulation, or im-
provements in brain function, or an eective immune sys-
tem. An example in plants would be the development
of the reproductive system in owering plants.
[44]
Such
adaptations may make the clade (monophyletic group)
more viable in a wide range of habitats. The acquisition
of such major adaptations has often served as the spark
for adaptive radiation, and huge success over long periods
of time for a whole group of animals or plants.
3.4 Compromise and conict between
adaptations
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.
[45]
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 oers a niche for
ectoparasites; the only ying penguins do is under water.
Adaptations serving dierent functions may be mutually
destructive. Compromise and makeshift occur widely,
not perfection. Selection pressures pull in dierent di-
rections, and the adaptation that results is some kind of
compromise.
[46]
Since the phenotype as a whole is the target
of selection, it is impossible to improve simul-
taneously all aspects of the phenotype to the
same degree. Ernst Mayr.
[47]
Consider the antlers of the Irish elk, (often supposed to
be far too large; in deer antler size has an allometric rela-
tionship 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.
[48]
Another ex-
ample: camouage to avoid detection is destroyed when
vivid colors are displayed at mating time. Here the risk to
life is counterbalanced by the necessity for reproduction.
Stream-dwelling salamanders, such as Caucasian Sala-
mander 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 pro-
tects their larvae frombeing washed out by current. How-
ever, elongated body increases risk of desiccation and de-
creases dispersal ability of the salamanders; it also nega-
tively aects their fecundity. As a result, re salamander,
less perfectly adapted to the mountain brook habitats, is
in general more successful, have a higher fecundity and
broader geographic range.
[49]
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 re-
duce his maneuverability and ight, and is hugely con-
spicuous; also, its growth costs food resources. Darwins
explanation of its advantage was in terms of sexual se-
lection: it depends on the advantage which certain indi-
viduals have over other individuals of the same sex and
6 5 RELATED ISSUES
species, in exclusive relation to reproduction.
[50]
The
kind of sexual selection represented by the peacock is
called 'mate choice', with an implication that the process
selects the more t over the less t, and so has survival
value.
[51]
The recognition of sexual selection was for a
long time in abeyance, but has been rehabilitated.
[52]
In
practice, the blue peafowl Pavo cristatus is a pretty suc-
cessful species, with a big natural range in India, so the
overall outcome of their mating system is quite viable.
The conict between the size of the human foetal brain
at birth, (which cannot be larger than about 400ccs, else
it will not get through the mothers pelvis) and the size
needed for an adult brain (about 1400ccs), means the
brain of a newborn child is quite immature. The most vi-
tal 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
fromour upright bipedal stance, without which our pelvis
could be shaped more suitably for birth. Neanderthals
had a similar problem.
[53][54][55]
As another example, the long neck of a girae is a burden
and a blessing. The necks of girae can be over eight feet
long. This neck can be used for inter-species competition
or for foraging on tall trees where shorter herbivores can-
not reach. However, as previously stated, there is always
a trade-o. This long neck is heavy and it adds to the
body mass of a girae, so the girae needs an abundance
of nutrition to provide for this costly adaptation.
[56]
4 Shifts in function
Adaptation and function are two aspects of
one problem. Julian Huxley
[57]
4.1 Pre-adaptations
This occurs when a species or population has character-
istics which (by chance) are suited for conditions which
have not yet arisen. For example, the polyploid rice-grass
Spartina townsendii is better adapted than either of its
parent species to their own habitat of saline marsh and
mud-ats.
[58]
White Leghorn fowl are markedly more re-
sistant to vitamin B deciency than other breeds.
[59]
On
a plentiful diet there is no dierence, but on a restricted
diet this preadaptation could be decisive.
Pre-adaptation may occur because a natural population
carries a huge quantity of genetic variability.
[60]
In diploid
eukaryotes, this is a consequence of the system of sexual
reproduction, where mutant alleles get partially shielded,
for example, by the selective advantage of heterozygotes.
Micro-organisms, with their huge populations, also carry
a great deal of genetic variability.
The rst experimental evidence of the pre-adaptive na-
ture of genetic variants in micro-organisms was provided
by Salvador Luria and Max Delbrck who developed
uctuation analysis, a method to show the random uc-
tuation of pre-existing genetic changes that conferred re-
sistance to phage in the bacterium Escherichia coli.
4.2 Co-option of existing traits: exapta-
tion
Main article: Exaptation
The classic example is the ear ossicles of mammals,
which we know from palaeontological and embryologi-
cal studies originated in the upper and lower jaws and the
hyoid of their Synapsid ancestors, and further back still
were part of the gill arches of early sh.
[61][62]
We owe
this esoteric knowledge to the comparative anatomists,
who, a century ago, were at the cutting edge of evolution-
ary studies.
[63]
The word exaptation was coined to cover
these shifts in function, which are surprisingly common in
evolutionary history.
[64]
The origin of wings from feath-
ers that were originally used for temperature regulation is
a more recent discovery (see feathered dinosaurs).
5 Related issues
5.1 Non-adaptive traits
Some traits do not appear to be adaptive, that is, they
appear to have a neutral or even deleterious eect on
tness in the current environment. Because genes have
pleiotropic eects, not all traits may be functional (i.e.
spandrels). Alternatively, a trait may have been adap-
tive at some point in an organisms evolutionary his-
tory, but a change in habitats caused what used to be an
adaptation to become unnecessary or even a hindrance
(maladaptations). Such adaptations are termed vestigial.
5.4 Flexibility, acclimatization, learning 7
5.1.1 Vestigial organs
Main article: Vestigiality
Many organisms have vestigial organs, which are the rem-
nants of fully functional structures in their ancestors. As
a result of changes in lifestyle the organs became redun-
dant, and are either not functional or reduced in function-
ality. With the loss of function goes the loss of positive
selection, and the subsequent accumulation of deleteri-
ous mutations. Since any structure represents some kind
of cost to the general economy of the body, an advan-
tage 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.
[65]
5.2 Fitness landscapes
Main article: Fitness landscape
Sewall Wright proposed that populations occupy adap-
tive peaks on a tness landscape. In order to evolve to an-
other, higher peak, a population would rst have to pass
through a valley of maladaptive intermediate stages.
[66]
A given population might be trapped on a peak that is
not optimally adapted.
5.3 Extinction
Main article: Extinction
If a population cannot move or change suciently to pre-
serve its long-term viability, then obviously, it will be-
come 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 (pop-
ulation, gene pool ...) exceeds the birth rate for a long
enough period for the species to disappear. It was an ob-
servation of Van Valen that groups of species tend to have
a characteristic and fairly regular rate of extinction.
[67]
5.3.1 Co-extinction
Main article: Co-extinction
Just as we have co-adaptation, there is also co-extinction.
Co-extinction refers to the loss of a species due to the
extinction of another; for example, the extinction of
parasitic insects following the loss of their hosts. Co-
extinction can also occur when a owering plant loses its
pollinator, or through the disruption of a food chain.
[68]
Species co-extinction is a manifestation of the intercon-
nectedness of organisms in complex ecosystems ... While
co-extinction may not be the most important cause of
species extinctions, it is certainly an insidious one.
[69]
5.4 Flexibility, acclimatization, learning
Flexibility deals with the relative capacity of an organ-
ism to maintain themselves in dierent habitats: their de-
gree of specialization. Acclimatization is a term used for
automatic physiological adjustments during life; learning
is the term used for improvement in behavioral perfor-
mance during life. In biology these terms are preferred,
not adaptation, for changes during life which improve the
performance of individuals. These adjustments are not
inherited genetically by the next generation.
Adaptation, on the other hand, occurs over many gener-
ations; it is a gradual process caused by natural selection
which changes the genetic make-up of a population so the
collective performs better in its niche.
5.4.1 Flexibility
Populations dier in their phenotypic plasticity, which is
the ability of an organismwith a given genotype to change
its phenotype in response to changes in its habitat, or to
move to a dierent habitat.
[70][71]
To a greater or lesser extent, all living things can adjust
to circumstances. The degree of exibility is inherited,
and varies to some extent between individuals. A highly
specialized animal or plant lives only in a well-dened
habitat, eats a specic type of food, and cannot survive if
its needs are not met. Many herbivores are like this; ex-
treme examples are koalas which depend on eucalyptus,
and pandas which require bamboo. A generalist, on the
other hand, eats a range of food, and can survive in many
dierent conditions. Examples are humans, rats, crabs
and many carnivores. The tendency to behave in a spe-
cialized or exploratory manner is inherited it is an adap-
tation.
Rather dierent is developmental exibility: An animal
8 6 FUNCTION AND TELEONOMY
or plant is developmentally exible if when it is raised or
transferred to new conditions it develops so that it is bet-
ter tted to survive in the new circumstances.
[72]
Once
again, there are huge dierences between species, and the
capacities to be exible are inherited.
5.4.2 Acclimatization
Main article: Acclimatization
If humans move to a higher altitude, respiration and phys-
ical exertion become a problem, but after spending time
in high altitude conditions they acclimatize to the pres-
sure by increasing production of red blood corpuscles.
The ability to acclimatize is an adaptation, but not the
acclimatization itself. Fecundity goes down, but deaths
from some tropical diseases also goes down.
Over a longer period of time, some people will repro-
duce better at these high altitudes than others. They will
contribute more heavily to later generations. Gradually
the whole population becomes adapted to the new con-
ditions. This we know takes place, because the perfor-
mance of long-term communities at higher altitude is sig-
nicantly better than the performance of new arrivals,
even when the new arrivals have had time to make phys-
iological adjustments.
[73]
Some kinds of acclimatization happen so rapidly that they
are better called reexes. The rapid colour changes in
some atsh, cephalopods, chameleons are examples.
[74]
5.4.3 Learning
Social learning is supreme for humans, and is possible for
quite a few mammals and birds: of course, that does not
involve genetic transmission except to the extent that the
capacities are inherited. Similarly, the capacity to learn
is an inherited adaptation, but not what is learnt; the ca-
pacity for human speech is inherited, but not the details
of language.
6 Function and teleonomy
Adaptation raises some issues concerning how biologists
use key terms such as function.
6.1 Function
To say something has a function is to say something about
what it does for the organism, obviously. It also says
something about its history: how it has come about. A
heart pumps blood: that is its function. It also emits
sound, which is just an ancillary side-eect. That is not its
function. The heart has a history (which may be well or
poorly understood), and that history is about how natural
selection formed and maintained the heart as a pump. Ev-
ery aspect of an organismthat has a function has a history.
Now, an adaptation must have a functional history: there-
fore we expect it must have undergone selection caused
by relative survival in its habitat. It would be quite wrong
to use the word adaptation about a trait which arose as a
by-product.
[75][76]
It is widely regarded as unprofessional for a biologist to
say something like A wing is for ying, although that
is their normal function. A biologist would be conscious
that sometime in the remote past feathers on a small di-
nosaur had the function of retaining heat, and that later
many wings were not used for ying (e.g. penguins,
ostriches). So, the biologist would rather say that the
wings on a bird or an insect usually had the function of
aiding ight. That would carry the connotation of being
an adaptation with a history of evolution by natural selec-
tion.
6.2 Teleonomy
Main article: Teleonomy
Teleonomy is a term invented to describe the study of
goal-directed functions which are not guided by the con-
scious forethought of man or any supernatural entity. It
is contrasted with Aristotle's teleology, which has conno-
tations of intention, purpose and foresight. Evolution is
teleonomic; adaptation hoards hindsight rather than fore-
sight. The following is a denition for its use in biology:
Teleonomy: The hypothesis that adaptations
arise without the existence of a prior purpose,
but by the action of natural selection on genetic
variability.
[77]
The term may have been suggested by Colin Pittendrigh
in 1958;
[78]
it grew out of cybernetics and self-organising
systems. Ernst Mayr, George C. Williams and Jacques
9
Monod picked up the term and used it in evolutionary
biology.
[79][80][81][82]
Philosophers of science have also commented on the
term. Ernest Nagel analysed the concept of goal-
directedness in biology;
[83]
and David Hull commented
on the use of teleology and teleonomy by biologists:
Haldane can be found remarking, Teleology
is like a mistress to a biologist: he cannot live
without her but hes unwilling to be seen with
her in public. Today the mistress has become
a lawfully wedded wife. Biologists no longer
feel obligated to apologize for their use of tele-
ological language; they aunt it. The only con-
cession which they make to its disreputable
past is to rename it teleonomy.
[84]
7 See also
Adaptive evolution in the human genome
Adaptive memory
Adaptive mutation
Adaptive radiation
Co-adaptation
Co-evolution
Ecological trap
Evolutionary physiology
Evolvability
Exaptation
Experimental evolution
Intragenomic conict
Mimicry
Neutral theory of molecular evolution
Phenotypic plasticity
Polymorphism (biology)
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Michael Hardy, Lexor, Gabbe, Habj, Big iron, Andres, Charles Matthews, Steinsky, DJ Clayworth, Samsara, Jerzy, Robbot, Sander123,
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Allens, NeilN, Elliskev, The Claw, SmackBot, Amcbride, Enlil Ninlil, ElectricRay, Gilliam, Skizzik, CanbekEsen, Chris the speller,
Unint, Miquonranger03, Ed Hagen, Royboycrashfan, NYKevin, Egsan Bacon, Dodgydobson, Liontooth, Addshore, EPM, LazyBoi633,
Ravenswood Media, Richard001, DMacks, Salamurai, GoldenTorc, SashatoBot, Mchavez, John, Heimstern, Joelmills, Extremophile,
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Epbr123, Headbomb, Marek69, SomeStranger, James086, Z10x, Dmitri Lytov, Nick Number, Pie Man 360, Mentisto, Porqin, Anti-
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Deathgod1995, Dr Dima, AntiSpamBot, Sewings, Juliancolton, SixteenBitJorge, Guyzero, Martial75, Idioma-bot, Je G., AlnoktaBOT,
Pilotbob, Philip Trueman, TXiKiBoT, Vipinhari, Vanished user ikijeirw34iuaeolaseric, Melsaran, JhsBot, Abdullais4u, Whatiguana,
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