Adaptation is the process that makes organisms better suited to their habitat. 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. 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. The following definitions are due to Theodosius Dobzhansky.

1. 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.

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. Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as a nutrient in a long-term laboratory experiment, Flavobacterium evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing, and the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic pesticide pentachlorophenol. 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).

Adaptation occurs through the gradual modification of existing structures. Consequently, structures with similar internal organisation 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. However, since all living organisms are related to some extent, 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.

During evolution, some structures may lose their original function and become vestigial structures. 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, the non-functional remains of eyes in blind cave-dwelling fish, wings in flightless birds, and the presence of hip bones in whales and snakes. Examples of vestigial structures in humans include wisdom teeth, the coccyx, the vermiform appendix, and other behavioural vestiges such as goose bumps 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. 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. Within cells, molecular machines such as the bacterial flagella and protein sorting machinery evolved by the recruitment of several pre-existing proteins that previously had different functions. 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.

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. 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 minimise the competition between them for food.

An area of current investigation in evolutionary developmental biology is the developmental basis of adaptations and exaptations. This research addresses the origin and evolution of embryonic development and how modifications of development and developmental processes produce novel features. These studies have shown that evolution can alter development to produce new structures, such as embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in mammals. 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. It is now becoming clear that most alterations in the form of organisms are due to changes in a small set of conserved genes.

Types of adaptations

"Adaptation is the heart and soul of evolution." -Niles Eldredge

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.

Habitat tracking

When a habitat changes, the most common thing to happen is that the resident population moves to another locale which suits it; this is the typical response of flying insects or oceanic organisms, who 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 thesis).

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 genetic variation that will lead to differing characteristics of offspring and hence abet adaptation. 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.[34] Genetic changes may result in visible structures, or may adjust physiological activity in a way that suits the changed habitat.

It is now clear that 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. Also, to a greater or lesser extent, the process affects every species in a particular ecosystem.

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 Queen's hypothesis.

Intimate relationships: co-adaptations

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 corresponding 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 flowering plants and insects (pollination).

The gut contents, wing structures, and mouthpart morphologies of fossilized beetles and flies suggest that they acted as early pollinators. The association between beetles and angiosperms during the early Cretaceous period led to parallel radiations of angiosperms and insects into the late Cretaceous. The evolution of nectaries in late Cretaceous flowers signals the beginning of the mutualism between hymenopterans and angiosperms.


Henry Walter 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. A common example seen in temperate gardens is the hover-fly, many of which – though bearing no sting – mimic the warning colouration of hymenoptera (wasps and bees). Such mimicry does not need to be perfect to improve the survival of the palatable species.


Bates, Wallace and Müller believed that Batesian and Müllerian mimicry provided evidence for the action of natural selection, a view which is now standard amongst biologists. All aspects of this situation can be, and have been, the subject of research. Field and experimental work on these ideas continues to this day; the topic connects strongly to speciation, genetics and development.

The basic machinery: internal adaptations

There are some important adaptations to do with the overall coordination of the systems in the body. Such adaptations may have significant consequences. Examples, in vertebrates, would be temperature regulation, or improvements in brain function, or an effective immune system. An example in plants would be the development of the reproductive system in flowering plants. 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.


Huxley, Julian 1942. Evolution the modern synthesis. Allen & Unwin, London. p449

Mayr, Ernst (1982). The growth of biological thought: diversity, evolution, and inheritance (1st ed.). Cambridge, Mass: Belknap Press. p. 483. ISBN 0-674-36445-7. "Adaptation... could no longer be considered a static condition, a product of a creative past, and became instead a continuing dynamic process."

Price P.W. 1980. The evolutionary biology of parasites. Princeton.

The Oxford Dictionary of Science defines adaptation as "Any change in the structure or functioning of an organism that makes it better suited to its environment".

Bowler, P.J. (2003) [1984]. Evolution: the history of an idea (3rd ed.). University of California Press. p. 10. ISBN 0-520-23693-9.

Patterson C. 1999. Evolution. Natural History Museum, London. p1

Williams, George C (1966). Adaptation and natural selection: a critique of some current evolutionary thought. Princeton University Press. p. 5. ISBN 0-691-02357-3. "Evolutionary adaptation is a phenomenon of pervasive importance in biology"

Mayr, Ernst (1963). Animal species and evolution (1st ed.). Cambridge: Belknap Press of Harvard University Press. ISBN 0-674-03750-2.

Mayr, Ernst (1982). The growth of biological thought: diversity, evolution, and inheritance (1st ed.). Cambridge, Mass: Belknap Press. pp. 562–566. ISBN 0-674-36445-7.

Salzburger W., Mack T., Verheyen E., Meyer A. (2005). "Out of Tanganyika: Genesis, explosive speciation, key-innovations and phylogeography of the haplochromine cichlid fishes" (PDF). BMC Evolutionary Biology 5: 17. doi:10.1186/1471-2148-5-17. PMC 554777. PMID 15723698.

Kornfield, Irv; Smith, Peter (November 2000). "African Cichlid Fishes: Model Systems for Evolutionary Biology". Annual Review of Ecology and Systematics 31: 163. doi:10.1146/annurev.ecolsys.31.1.163.

Stebbins, G. Ledyard, Jr. 1950. Variation and evolution in plants. Columbia. Polyploidy, chapters 8 and 9.

Margulis, Lynn (ed) 1991. Symbiosis as a source of evolutionary innovation: speciation and morphogenesis MIT. ISBN 0-262-13269-9

Hutchinson, G. Evelyn (1965). The ecological theatre and the evolutionary play. Yale. ISBN 0-300-00586-5. The niche is the central concept in evolutionary ecology; see especially part II The niche: an abstractly inhabited hypervolume. p26–78

Dobzhansky, T.; Hecht, MK; Steere, WC (1968). "On some fundamental concepts of evolutionary biology". Evolutionary biology volume 2 (1st ed.). New York: Appleton-Century-Crofts. pp. 1–34.

Dobzhansky, T. (1970). Genetics of the evolutionary process. N.Y.: Columbia. pp. 4–6, 79–82, 84–87. ISBN 0-231-02837-7.

Dobzhansky T. (1956). "Genetics of natural populations XXV. Genetic changes in populations of Drosophila pseudoobscura and Drosphila persimilis in some locations in California". Evolution 10 (1): 82–92. doi:10.2307/2406099. JSTOR 2406099.

Endler, John A (1986). "Fitness and adaptation". Natural selection in the wild. Princeton University Press. pp. 33–51. ISBN 0-691-08387-8.

Sober, Elliott (1984). The nature of selection: evolutionary theory in philosophical focus. Cambridge, Mass: MIT Press. ISBN 0-262-19232-2.

Futuyma D.J. 1986. Evolution. 2nd ed, Sinauer, Sunderland, Massachusetts.

Desmond, Adrian 1989. The politics of evolution. Chicago. p31/32, footnote 18.

Sober, Elliott 1993. Philosophy of biology. Oxford. Chapter 2

Darwin, Charles. 1872. The origin of species. 6th edition, p397: Rudimentary, atrophied and aborted organs.

Peter H. 2003. Evolution: the history of an idea. 3rd ed, California. p86–95, especially "Whatever the true nature of Lamark's theory, it was his mechanism of adaptation that caught the attention of later naturalists". (p90)

Provine, William 1986. Sewall Wright and evolutionary biology. University of Chicago Press.

Ford E.B. 1975. Ecological genetics, 4th ed. Chapman and Hall, London.

Orr H. (2005). "The genetic theory of adaptation: a brief history". Nature Rev. Genetics 6 (2): 119–127. doi:10.1038/nrg1523. PMID 15716908.

Eldredge, Niles 1995. Reinventing Darwin: the great evolutionary debate. Wiley N.Y. p33

Eldredge, Niles 1986. Time frames: the rethinking of Darwinian evolution and the theory of punctuated equilibria. p136, Of glaciers and beetles.

Eldredge, Niles 1995. Reinventing Darwin: the great evolutionary debate. Wiley, N.Y. p64

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