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Adaptation (disambiguation)

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116-1499: Adaptation , in biology, is the process or trait by which organisms or population better match their environment Adaptation may also refer to: Adaptation Collective intelligence Collective action Self-organized criticality Herd mentality Phase transition Agent-based modelling Synchronization Ant colony optimization Particle swarm optimization Swarm behaviour Social network analysis Small-world networks Centrality Motifs Graph theory Scaling Robustness Systems biology Dynamic networks Evolutionary computation Genetic algorithms Genetic programming Artificial life Machine learning Evolutionary developmental biology Artificial intelligence Evolutionary robotics Reaction–diffusion systems Partial differential equations Dissipative structures Percolation Cellular automata Spatial ecology Self-replication Conversation theory Entropy Feedback Goal-oriented Homeostasis Information theory Operationalization Second-order cybernetics Self-reference System dynamics Systems science Systems thinking Sensemaking Variety Ordinary differential equations Phase space Attractors Population dynamics Chaos Multistability Bifurcation Rational choice theory Bounded rationality In biology , adaptation has three related meanings. Firstly, it

232-484: 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 . 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 "

348-451: A genealogical branching of a single evolutionary tree , in which "It is absurd to talk of one animal being higher than another", thereby discarding Lamarck's idea of independent lineages progressing to higher forms. While developing this intensive study of transmutation, Darwin became mired in more work. Still rewriting his Journal , he took on editing and publishing the expert reports on his collections, and with Henslow's help obtained

464-441: 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

580-562: A 230-page "Essay", to be expanded with his research results if he died prematurely. In November, the anonymously published sensational best-seller Vestiges of the Natural History of Creation brought wide interest in transmutation. Darwin scorned its amateurish geology and zoology, but carefully reviewed his own arguments. Controversy erupted, and it continued to sell well despite contemptuous dismissal by scientists. Darwin completed his third geological book in 1846. He now renewed

696-845: A Bachelor of Arts degree as the first step towards becoming an Anglican country parson . Darwin was unqualified for Cambridge's Tripos exams and was required instead to join the ordinary degree course. He preferred riding and shooting to studying. During the first few months of Darwin's enrolment at Christ's College, his second cousin William Darwin Fox was still studying there. Fox impressed him with his butterfly collection, introducing Darwin to entomology and influencing him to pursue beetle collecting. He did this zealously and had some of his finds published in James Francis Stephens ' Illustrations of British entomology (1829–1932). Through Fox, Darwin became

812-673: A Treasury grant of £1,000 to sponsor this multi-volume Zoology of the Voyage of H.M.S. Beagle , a sum equivalent to about £115,000 in 2021. He stretched the funding to include his planned books on geology, and agreed to unrealistic dates with the publisher. As the Victorian era began, Darwin pressed on with writing his Journal , and in August 1837 began correcting printer's proofs . As Darwin worked under pressure, his health suffered. On 20 September, he had "an uncomfortable palpitation of

928-591: A beginning to each crop of species". Darwin's book The Structure and Distribution of Coral Reefs on his theory of atoll formation was published in May 1842 after more than three years of work, and he then wrote his first "pencil sketch" of his theory of natural selection. To escape the pressures of London, the family moved to rural Down House in Kent in September. On 11 January 1844, Darwin mentioned his theorising to

1044-547: A book on geology. When they reached Brazil, Darwin was delighted by the tropical forest , but detested the sight of slavery there , and disputed this issue with FitzRoy. The survey continued to the south in Patagonia . They stopped at Bahía Blanca , and in cliffs near Punta Alta Darwin made a major find of fossil bones of huge extinct mammals beside modern seashells, indicating recent extinction with no signs of change in climate or catastrophe. He found bony plates like

1160-718: A break and went "geologising" in Scotland. He visited Glen Roy in glorious weather to see the parallel "roads" cut into the hillsides at three heights. He later published his view that these were marine-raised beaches, but then had to accept that they were shorelines of a proglacial lake . Fully recuperated, he returned to Shrewsbury in July 1838. Used to jotting down daily notes on animal breeding, he scrawled rambling thoughts about marriage, career and prospects on two scraps of paper, one with columns headed "Marry" and "Not Marry" . Advantages under "Marry" included "constant companion and

1276-471: A calm spell. On their first stop ashore at St Jago in Cape Verde, Darwin found that a white band high in the volcanic rock cliffs included seashells. FitzRoy had given him the first volume of Charles Lyell 's Principles of Geology , which set out uniformitarian concepts of land slowly rising or falling over immense periods, and Darwin saw things Lyell's way, theorising and thinking of writing

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1392-562: A close friend and follower of botany professor John Stevens Henslow . He met other leading parson-naturalists who saw scientific work as religious natural theology , becoming known to these dons as "the man who walks with Henslow". When his own exams drew near, Darwin applied himself to his studies and was delighted by the language and logic of William Paley 's Evidences of Christianity (1795). In his final examination in January 1831, Darwin did well, coming tenth out of 178 candidates for

1508-412: A copy of his journal for his family. He had some expertise in geology, beetle collecting and dissecting marine invertebrates, but in all other areas, was a novice and ably collected specimens for expert appraisal. Despite suffering badly from seasickness, Darwin wrote copious notes while on board the ship. Most of his zoology notes are about marine invertebrates, starting with plankton collected during

1624-596: A fascination and expertise in marine invertebrates, dating back to his student days with Grant, by dissecting and classifying the barnacles he had collected on the voyage, enjoying observing beautiful structures and thinking about comparisons with allied structures. In 1847, Hooker read the "Essay" and sent notes that provided Darwin with the calm critical feedback that he needed, but would not commit himself and questioned Darwin's opposition to continuing acts of creation . In an attempt to improve his chronic ill health, Darwin went in 1849 to Dr. James Gully 's Malvern spa and

1740-529: A fortnight mapping strata in Wales. After leaving Sedgwick in Wales, Darwin spent a few days with student friends at Barmouth . He returned home on 29 August to find a letter from Henslow proposing him as a suitable (if unfinished) naturalist for a self-funded supernumerary place on HMS  Beagle with captain Robert FitzRoy , a position for a gentleman rather than "a mere collector". The ship

1856-532: A freed black slave who had accompanied Charles Waterton in the South American rainforest . In Darwin's second year at the university, he joined the Plinian Society , a student natural-history group featuring lively debates in which radical democratic students with materialistic views challenged orthodox religious concepts of science. He assisted Robert Edmond Grant 's investigations of

1972-473: A friend in old age ... better than a dog anyhow", against points such as "less money for books" and "terrible loss of time". Having decided in favour of marriage, he discussed it with his father, then went to visit his cousin Emma on 29 July. At this time he did not get around to proposing, but against his father's advice, he mentioned his ideas on transmutation. He married Emma on 29 January 1839 and they were

2088-530: A giant capybara . The armour fragments were actually from Glyptodon , a huge armadillo-like creature, as Darwin had initially thought. These extinct creatures were related to living species in South America. In mid-December, Darwin took lodgings in Cambridge to arrange expert classification of his collections, and prepare his own research for publication. Questions of how to combine his diary into

2204-556: A giant version of the armour on local armadillos . From a jaw and tooth he identified the gigantic Megatherium , then from Cuvier's description thought the armour was from this animal. The finds were shipped to England, and scientists found the fossils of great interest. In Patagonia, Darwin came to wrongly believe the territory was devoid of reptiles. On rides with gauchos into the interior to explore geology and collect more fossils, Darwin gained social, political and anthropological insights into both native and colonial people at

2320-526: 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 . The word is controversial because it

2436-506: A great name among the Naturalists of Europe". On 2 October 1836, Beagle anchored at Falmouth , Cornwall. Darwin promptly made the long coach journey to Shrewsbury to visit his home and see relatives. He then hurried to Cambridge to see Henslow, who advised him on finding available naturalists to catalogue Darwin's animal collections and to take on the botanical specimens. Darwin's father organised investments, enabling his son to be

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2552-427: A higher fecundity and broader geographic range. 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

2668-410: A majority of the educated public had accepted evolution as a fact . However, many initially favoured competing explanations that gave only a minor role to natural selection, and it was not until the emergence of the modern evolutionary synthesis from the 1930s to the 1950s that a broad consensus developed in which natural selection was the basic mechanism of evolution. Darwin's scientific discovery

2784-423: 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. The significance of an adaptation can only be understood in relation to

2900-457: 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. It is a profound truth that Nature does not know best; that genetical evolution... is a story of waste, makeshift, compromise and blunder. Since

3016-632: A poetic fantasy of gradual creation including undeveloped ideas anticipating concepts his grandson expanded. Both families were largely Unitarian , though the Wedgwoods were adopting Anglicanism . Robert Darwin, a freethinker , had baby Charles baptised in November 1809 in the Anglican St Chad's Church, Shrewsbury , but Charles and his siblings attended the local Unitarian Church with their mother. The eight-year-old Charles already had

3132-609: A process he called natural selection , in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding . Darwin has been described as one of the most influential figures in human history and was honoured by burial in Westminster Abbey . Darwin's early interest in nature led him to neglect his medical education at the University of Edinburgh ; instead, he helped to investigate marine invertebrates . His studies at

3248-539: A programmer of laws. Darwin stayed with his freethinking brother Erasmus, part of this Whig circle and a close friend of the writer Harriet Martineau , who promoted the Malthusianism that underpinned the controversial Whig Poor Law reforms to stop welfare from causing overpopulation and more poverty. As a Unitarian, she welcomed the radical implications of transmutation of species , promoted by Grant and younger surgeons influenced by Geoffroy . Transmutation

3364-591: A sand beach. He theorised that as the land rose, oceanic islands sank, and coral reefs round them grew to form atolls . On the geologically new Galápagos Islands , Darwin looked for evidence attaching wildlife to an older "centre of creation", and found mockingbirds allied to those in Chile but differing from island to island. He heard that slight variations in the shape of tortoise shells showed which island they came from, but failed to collect them, even after eating tortoises taken on board as food. In Australia,

3480-517: A self-funded gentleman scientist , and an excited Darwin went around the London institutions being fêted and seeking experts to describe the collections. British zoologists at the time had a huge backlog of work, due to natural history collecting being encouraged throughout the British Empire, and there was a danger of specimens just being left in storage. Charles Lyell eagerly met Darwin for

3596-426: A similar problem. 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. 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

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3712-719: A similarity between farmers picking the best stock in selective breeding, and a Malthusian Nature selecting from chance variants so that "every part of newly acquired structure is fully practical and perfected", thinking this comparison "a beautiful part of my theory". He later called his theory natural selection , an analogy with what he termed the "artificial selection" of selective breeding. On 11 November, he returned to Maer and proposed to Emma, once more telling her his ideas. She accepted, then in exchanges of loving letters she showed how she valued his openness in sharing their differences, while expressing her strong Unitarian beliefs and concerns that his honest doubts might separate them in

3828-705: A taste for natural history and collecting when he joined the day school run by its preacher in 1817. That July, his mother died. From September 1818, he joined his older brother Erasmus in attending the nearby Anglican Shrewsbury School as a boarder . Darwin spent the summer of 1825 as an apprentice doctor, helping his father treat the poor of Shropshire, before going to the well-regarded University of Edinburgh Medical School with his brother Erasmus in October 1825. Darwin found lectures dull and surgery distressing, so he neglected his studies. He learned taxidermy in around 40 daily hour-long sessions from John Edmonstone ,

3944-468: A time of revolution, and learnt that two types of rhea had separate but overlapping territories. Further south, he saw stepped plains of shingle and seashells as raised beaches at a series of elevations. He read Lyell's second volume and accepted its view of "centres of creation" of species, but his discoveries and theorising challenged Lyell's ideas of smooth continuity and of extinction of species. Three Fuegians on board, who had been seized during

4060-411: A trait, whereas fitness predicts a trait's future. 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. Adaptation is the heart and soul of evolution. Before Darwin, adaptation

4176-598: 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. 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. Physiological resistance to the heart poisons ( cardiac glycosides ) that monarch butterflies store in their bodies to protect themselves from predators are driven by adaptive mutations in

4292-424: 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. 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

4408-523: 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. 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

4524-408: Is better fitted to survive in the new environment," writes the evolutionary biologist John Maynard Smith . 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

4640-558: Is called habitat tracking . It is one explanation put forward for the periods of apparent stasis in the fossil record (the punctuated equilibrium theory). 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. One example

4756-471: Is known as a Malthusian catastrophe . Darwin was well-prepared to compare this to Augustin de Candolle 's "warring of the species" of plants and the struggle for existence among wildlife, explaining how numbers of a species kept roughly stable. As species always breed beyond available resources, favourable variations would make organisms better at surviving and passing the variations on to their offspring, while unfavourable variations would be lost. He wrote that

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4872-506: 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. Genetic changes may result in entirely new or gradual change to visible structures, or they may adjust physiological activity in

4988-440: 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

5104-522: 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 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,

5220-766: 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

5336-571: Is the unifying theory of the life sciences , explaining the diversity of life . Darwin was born in Shrewsbury , Shropshire, on 12 February 1809, at his family's home, The Mount . He was the fifth of six children of wealthy society doctor and financier Robert Darwin and Susannah Darwin (née Wedgwood). His grandfathers Erasmus Darwin and Josiah Wedgwood were both prominent abolitionists . Erasmus Darwin had praised general concepts of evolution and common descent in his Zoonomia (1794),

5452-409: 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

5568-464: 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

5684-554: The Narrative were resolved at the end of the month when FitzRoy accepted Broderip's advice to make it a separate volume, and Darwin began work on his Journal and Remarks . Darwin's first paper showed that the South American landmass was slowly rising, and with Lyell's enthusiastic backing he read it to the Geological Society of London on 4 January 1837. On the same day, he presented his mammal and bird specimens to

5800-610: The Royal Society 's Royal Medal, and it made his reputation as a biologist . Upon the conclusion of his research, Darwin declared "I hate a barnacle as no man ever did before." In 1854, he became a Fellow of the Linnean Society of London , gaining postal access to its library. He began a major reassessment of his theory of species, and in November realised that divergence in the character of descendants could be explained by them becoming adapted to "diversified places in

5916-410: The University of Cambridge 's Christ's College from 1828 to 1831 encouraged his passion for natural science . However, it was his five-year voyage on HMS  Beagle from 1831 to 1836 that truly established Darwin as an eminent geologist. The observations and theories he developed during his voyage supported Charles Lyell 's concept of gradual geological change . Publication of his journal of

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6032-609: The White Leghorn chicken is markedly more resistant to vitamin B 1 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

6148-942: The Zoological Society . The ornithologist John Gould soon announced that the Galápagos birds that Darwin had thought a mixture of blackbirds , " gros-beaks " and finches , were, in fact, twelve separate species of finches . On 17 February, Darwin was elected to the Council of the Geological Society, and Lyell's presidential address presented Owen's findings on Darwin's fossils, stressing geographical continuity of species as supporting his uniformitarian ideas. Early in March, Darwin moved to London to be near this work, joining Lyell's social circle of scientists and experts such as Charles Babbage , who described God as

6264-521: The first Beagle voyage then given Christian education in England, were returning with a missionary. Darwin found them friendly and civilised, yet at Tierra del Fuego he met "miserable, degraded savages", as different as wild from domesticated animals. He remained convinced that, despite this diversity, all humans were interrelated with a shared origin and potential for improvement towards civilisation. Unlike his scientist friends, he now thought there

6380-492: 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

6496-412: 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

6612-519: The marsupial rat-kangaroo and the platypus seemed so unusual that Darwin thought it was almost as though two distinct Creators had been at work. He found the Aborigines "good-humoured & pleasant", their numbers depleted by European settlement. FitzRoy investigated how the atolls of the Cocos (Keeling) Islands had formed, and the survey supported Darwin's theorising. FitzRoy began writing

6728-598: The ordinary degree. Darwin had to stay at Cambridge until June 1831. He studied Paley's Natural Theology or Evidences of the Existence and Attributes of the Deity (first published in 1802), which made an argument for divine design in nature , explaining adaptation as God acting through laws of nature . He read John Herschel 's new book, Preliminary Discourse on the Study of Natural Philosophy (1831), which described

6844-518: 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. 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. All adaptations have a downside: horse legs are great for running on grass, but they cannot scratch their backs; mammals ' hair helps temperature, but offers

6960-596: The "final cause of all this wedging, must be to sort out proper structure, & adapt it to changes", so that "One may say there is a force like a hundred thousand wedges trying force into every kind of adapted structure into the gaps of in the economy of nature, or rather forming gaps by thrusting out weaker ones." This would result in the formation of new species. As he later wrote in his Autobiography : In October 1838, that is, fifteen months after I had begun my systematic enquiry, I happened to read for amusement Malthus on Population, and being well prepared to appreciate

7076-622: The Emotions in Man and Animals (1872). His research on plants was published in a series of books, and in his final book, The Formation of Vegetable Mould, through the Actions of Worms (1881), he examined earthworms and their effect on soil. Darwin published his theory of evolution with compelling evidence in his 1859 book On the Origin of Species . By the 1870s, the scientific community and

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7192-575: The ability of an organism with a given genotype (genetic type) to change its phenotype (observable characteristics) 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

7308-402: 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

7424-459: The afterlife. While he was house-hunting in London, bouts of illness continued and Emma wrote urging him to get some rest, almost prophetically remarking "So don't be ill any more my dear Charley till I can be with you to nurse you." He found what they called "Macaw Cottage" (because of its gaudy interiors) in Gower Street , then moved his "museum" in over Christmas. On 24 January 1839, Darwin

7540-506: The anatomy and life cycle of marine invertebrates in the Firth of Forth , and on 27 March 1827 presented at the Plinian his own discovery that black spores found in oyster shells were the eggs of a skate leech . One day, Grant praised Lamarck 's evolutionary ideas . Darwin was astonished by Grant's audacity, but had recently read similar ideas in his grandfather Erasmus' journals. Darwin

7656-718: The animals have modified. 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. Charles Darwin Charles Robert Darwin ( / ˈ d ɑːr w ɪ n / DAR -win ; 12 February 1809 – 19 April 1882)

7772-427: 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. Bates' work on Amazonian butterflies led him to develop

7888-401: 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

8004-525: The best of all possible worlds ." Voltaire 's satire Dr. Pangloss is a parody of this optimistic idea, and David Hume also argued against design. Charles Darwin broke with the tradition by emphasising the flaws and limitations which occurred in the animal and plant worlds. 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

8120-441: 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 . An example widely used today to study the interplay of adaptation and speciation

8236-437: The botanist Joseph Dalton Hooker , writing with melodramatic humour "it is like confessing a murder". Hooker replied, "There may, in my opinion, have been a series of productions on different spots, & also a gradual change of species. I shall be delighted to hear how you think that this change may have taken place, as no presently conceived opinions satisfy me on the subject." By July, Darwin had expanded his "sketch" into

8352-550: The brink of extinction in a process called evolutionary rescue . Adaptation does affect, to some extent, every species in a particular ecosystem . 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. Existing genetic variation and mutation were

8468-542: The duties of Secretary of the Geological Society. After initially declining the work, he accepted the post in March 1838. Despite the grind of writing and editing the Beagle reports, Darwin made remarkable progress on transmutation, taking every opportunity to question expert naturalists and, unconventionally, people with practical experience in selective breeding such as farmers and pigeon fanciers . Over time, his research drew on information from his relatives and children,

8584-477: 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

8700-713: The family butler, neighbours, colonists and former shipmates. He included mankind in his speculations from the outset, and on seeing an orangutan in the zoo on 28 March 1838 noted its childlike behaviour. The strain took a toll, and by June he was being laid up for days on end with stomach problems, headaches and heart symptoms. For the rest of his life, he was repeatedly incapacitated with episodes of stomach pains, vomiting, severe boils , palpitations, trembling and other symptoms, particularly during times of stress, such as attending meetings or making social visits. The cause of Darwin's illness remained unknown, and attempts at treatment had only ephemeral success. On 23 June, he took

8816-465: 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 (the model), 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

8932-597: The first time on 29 October and soon introduced him to the up-and-coming anatomist Richard Owen , who had the facilities of the Royal College of Surgeons to work on the fossil bones collected by Darwin. Owen's surprising results included other gigantic extinct ground sloths as well as the Megatherium Darwin had identified, a near complete skeleton of the unknown Scelidotherium and a hippopotamus -sized rodent -like skull named Toxodon resembling

9048-582: The heart", so his doctors urged him to "knock off all work" and live in the country for a few weeks. After visiting Shrewsbury, he joined his Wedgwood relatives at Maer Hall , Staffordshire, but found them too eager for tales of his travels to give him much rest. His charming, intelligent, and cultured cousin Emma Wedgwood , nine months older than Darwin, was nursing his invalid aunt. His uncle Josiah pointed out an area of ground where cinders had disappeared under loam and suggested that this might have been

9164-472: The highest aim of natural philosophy as understanding such laws through inductive reasoning based on observation, and Alexander von Humboldt 's Personal Narrative of scientific travels in 1799–1804. Inspired with "a burning zeal" to contribute, Darwin planned to visit Tenerife with some classmates after graduation to study natural history in the tropics. In preparation, he joined Adam Sedgwick 's geology course, then on 4 August travelled with him to spend

9280-579: The life of an organism. The following definitions are given by the evolutionary biologist Theodosius Dobzhansky : 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; learning means alteration in behavioural performance during life. Flexibility stems from phenotypic plasticity ,

9396-433: The mother's pelvis ) and the size needed for an adult brain (about 1400 cm ), 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

9512-429: The neck and to carry its weight around. Adaptation and function are two aspects of one problem. 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,

9628-411: 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. 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

9744-433: The official Narrative of the Beagle voyages, and after reading Darwin's diary, he proposed incorporating it into the account. Darwin's Journal was eventually rewritten as a separate third volume, on geology and natural history. In Cape Town , South Africa, Darwin and FitzRoy met John Herschel, who had recently written to Lyell praising his uniformitarianism as opening bold speculation on "that mystery of mysteries,

9860-496: The origin of species". Without telling Darwin, extracts from his letters to Henslow had been read to scientific societies, printed as a pamphlet for private distribution among members of the Cambridge Philosophical Society , and reported in magazines, including The Athenaeum . Darwin first heard of this at Cape Town, and at Ascension Island read of Sedgwick's prediction that Darwin "will have

9976-423: 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

10092-482: The other species, such as with flowering plants and pollinating insects . In mimicry , species evolve to resemble other species; in 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

10208-522: The parents of ten children, seven of whom survived to adulthood. Continuing his research in London, Darwin's wide reading now included the sixth edition of Malthus 's An Essay on the Principle of Population . On 28 September 1838, he noted its assertion that human "population, when unchecked, goes on doubling itself every twenty-five years, or increases in a geometrical ratio", a geometric progression so that population soon exceeds food supply in what

10324-507: 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. 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). 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

10440-573: The possibility that "one species does change into another" to explain the geographical distribution of living species such as the rheas, and extinct ones such as the strange extinct mammal Macrauchenia , which resembled a giant guanaco , a llama relative. Around mid-July, he recorded in his "B" notebook his thoughts on lifespan and variation across generations – explaining the variations he had observed in Galápagos tortoises , mockingbirds, and rheas. He sketched branching descent, and then

10556-407: The process of adaptation is never fully complete. 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

10672-471: 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

10788-520: The replacement of extinct species by others" as "a natural in contradistinction to a miraculous process". When organising his notes as the ship sailed home, Darwin wrote that, if his growing suspicions about the mockingbirds, the tortoises and the Falkland Islands fox were correct, "such facts undermine the stability of Species", then cautiously added "would" before "undermine". He later wrote that such facts "seemed to me to throw some light on

10904-519: The same idea, prompting the immediate joint submission of both their theories to the Linnean Society of London . Darwin's work established evolutionary descent with modification as the dominant scientific explanation of natural diversification. In 1871, he examined human evolution and sexual selection in The Descent of Man, and Selection in Relation to Sex , followed by The Expression of

11020-500: 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 cm , else it will not get through

11136-526: 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

11252-408: The struggle for existence which everywhere goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here, then, I had at last got a theory by which to work... By mid-December, Darwin saw

11368-409: The systematics of the taxa. He wrote his first examination of the species in 1846, but did not formally describe it until 1854. FitzRoy's long delayed Narrative was published in May 1839. Darwin's Journal and Remarks got good reviews as the third volume, and on 15 August it was published on its own. Early in 1842, Darwin wrote about his ideas to Charles Lyell, who noted that his ally "denies seeing

11484-569: The target of the poison, the sodium pump , resulting in target site insensitivity. 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. Convergence at the gene-level across distantly related species can arise because of evolutionary constraint. Habitats and biota do frequently change over time and space. Therefore, it follows that

11600-535: 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 and Alfred Russel Wallace 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

11716-458: The total biology of the species. 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

11832-468: 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 . 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

11948-603: 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. 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

12064-473: The voyage made Darwin famous as a popular author. Puzzled by the geographical distribution of wildlife and fossils he collected on the voyage, Darwin began detailed investigations and, in 1838, devised his theory of natural selection. Although he discussed his ideas with several naturalists, he needed time for extensive research, and his geological work had priority. He was writing up his theory in 1858 when Alfred Russel Wallace sent him an essay that described

12180-420: The voyage began on 27 December 1831; it lasted almost five years. As FitzRoy had intended, Darwin spent most of that time on land investigating geology and making natural history collections, while HMS Beagle surveyed and charted coasts. He kept careful notes of his observations and theoretical speculations, and at intervals during the voyage his specimens were sent to Cambridge together with letters including

12296-403: The work of earthworms , inspiring "a new & important theory" on their role in soil formation , which Darwin presented at the Geological Society on 1 November 1837. His Journal was printed and ready for publication by the end of February 1838, as was the first volume of the Narrative , but FitzRoy was still working hard to finish his own volume. William Whewell pushed Darwin to take on

12412-707: Was elected a Fellow of the Royal Society (FRS). On 29 January, Darwin and Emma Wedgwood were married at Maer in an Anglican ceremony arranged to suit the Unitarians, then immediately caught the train to London and their new home. Darwin now had the framework of his theory of natural selection "by which to work", as his "prime hobby". His research included extensive experimental selective breeding of plants and animals, finding evidence that species were not fixed and investigating many detailed ideas to refine and substantiate his theory. For fifteen years this work

12528-418: Was a " wren " was in the finch group . Darwin had not labelled the finches by island, but from the notes of others on the ship, including FitzRoy, he allocated species to islands. The two rheas were distinct species, and on 14 March Darwin announced how their distribution changed going southwards. By mid-March 1837, barely six months after his return to England, Darwin was speculating in his Red Notebook on

12644-426: Was an English naturalist , geologist , and biologist , widely known for his contributions to evolutionary biology . His proposition that all species of life have descended from a common ancestor is now generally accepted and considered a fundamental scientific concept. In a joint publication with Alfred Russel Wallace , he introduced his scientific theory that this branching pattern of evolution resulted from

12760-472: Was anathema to Anglicans defending social order, but reputable scientists openly discussed the subject, and there was wide interest in John Herschel's letter praising Lyell's approach as a way to find a natural cause of the origin of new species. Gould met Darwin and told him that the Galápagos mockingbirds from different islands were separate species, not just varieties, and what Darwin had thought

12876-425: Was in the background to his main occupation of writing on geology and publishing expert reports on the Beagle collections, in particular, the barnacles . The impetus of Darwin's barnacle research came from a collection of a barnacle colony from Chile in 1835, which he dubbed Mr. Arthrobalanus . His confusion over the relationship of this species ( Cryptophialus minutus ) to other barnacles caused him to fixate on

12992-504: Was no unbridgeable gap between humans and animals. A year on, the mission had been abandoned. The Fuegian they had named Jemmy Button lived like the other natives, had a wife, and had no wish to return to England. Darwin experienced an earthquake in Chile in 1835 and saw signs that the land had just been raised, including mussel -beds stranded above high tide. High in the Andes he saw seashells, and several fossil trees that had grown on

13108-577: Was rather bored by Robert Jameson 's natural-history course, which covered geology – including the debate between neptunism and plutonism . He learned the classification of plants and assisted with work on the collections of the University Museum , one of the largest museums in Europe at the time. Darwin's neglect of medical studies annoyed his father, who sent him to Christ's College, Cambridge , in January 1828, to study for

13224-414: 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

13340-574: Was surprised to find some benefit from hydrotherapy . Then, in 1851, his treasured daughter Annie fell ill, reawakening his fears that his illness might be hereditary. She died the same year after a long series of crises. In eight years of work on barnacles, Darwin's theory helped him to find " homologies " showing that slightly changed body parts served different functions to meet new conditions, and in some genera he found minute males parasitic on hermaphrodites , showing an intermediate stage in evolution of distinct sexes . In 1853, it earned him

13456-446: Was to leave in four weeks on an expedition to chart the coastline of South America. Robert Darwin objected to his son's planned two-year voyage, regarding it as a waste of time, but was persuaded by his brother-in-law, Josiah Wedgwood II , to agree to (and fund) his son's participation. Darwin took care to remain in a private capacity to retain control over his collection, intending it for a major scientific institution. After delays,

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