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It seems certain that the first "fiction" film, L'arroseur arrosé (The Waterer Watered, 1895) by Louis Lumière (1864–1948), was based on an 1889 comic strip by "Christophe" and that two of the most famous early American narrative films, Edwin S. Porter's (1869–1941) The Great Train Robbery (1903) and Dream of a Rarebit Fiend (1906), were derived, at least in part, from contemporary theatrical and comic strip material respectively. Generally the earliest attempts at narrative cinema were taken from already existing literary or theatrical sources and have provided by far the largest proportion of script material for the cinema ever since. This process, however, has been regularly plagued by arguments over the vexed question of fidelity. To what extent should (or can) a film be "faithful" to its original source? Which aspects of literary or theatrical technique are compatible with the film medium and which cannot be successfully transferred? To what extent should filmmakers alter characterization, setting, or plot to suit their own interpretation of the original? Does it matter if the filmmaker changes the original almost completely and yet comes up with a cinematic masterpiece in its own right? Should a film adaptation, in other words, always have to justify itself in terms of its closeness to its literary original, or can the two be accepted and judged independently?

The questions continue to be debated. Most theorizing tends to split types of adaptation into three categories: strict, loose, or free (using these or somewhat similar terms). They also often distinguish between classic or well-known works where audiences already have some knowledge of the original and may expect to see this reproduced reasonably faithfully on the screen, and less famous or forgotten works where audience loyalty to the original is less significant. Many critics accept a compromise: if the essence of the original (theme, mood, tone in particular) is preserved and not deliberately or incompetently distorted, then other, less crucial, changes are acceptable. The claim that a successful adaptation should be medium specific—thoroughly rethought in terms of film and the filmmaker's own creative approach and not hampered by inappropriate adherence to literary or stage techniques—is also now commonly held. Such a view, for example, would approve of A Clockwork Orange (1971) by Stanley Kubrick (1928–1999), despite its being disowned by the author of the original novel, Anthony Burgess (1917–1993), who felt that Kubrick overemphasized the violent and negative aspects of the book.

The most difficult task for the filmmaker is probably to take a classic or currently popular work and present it in a way that avoids alienating those who have a commitment to their own interpretation of the original while simultaneously producing something that works successfully as a film in its own right. These adaptations would normally fall into the category of strict or loose, though free reworkings of, for example, William Shakespeare (1564–1616) (Joe MacBeth, 1955), Charles Dickens (1812–1870) (Rich's Man's Folly, 1931; based on Dombey and Son), or Jane Austen (1775–1817) (Clueless, 1995; based on Emma) certainly exist. One of the most highly acclaimed examples of an adaptation that has managed to please both die-hard admirers of the original books and to be accepted as a cinematic masterpiece is Peter Jackson's (b. 1961) version of J. R. R. Tolkien's (1892–1973) The Lord of the Rings trilogy (2001–2003).

A more common resource, however, has been to take works that, for reasons of literary style, plot, or characterization, are more amenable to being "tampered with" and are less complete or self-sufficient in their original form, or that belong to literary genres such as detective or gangster fiction, thrillers, westerns, or science fiction, which are often considered to be marginal in terms of literary respectability and are thus less likely to arouse indignation if they are "betrayed" in the process of adaptation. Many of the finest American films fall into these categories, as do those of the French New Wave works that were based on Série noire (1979) or pulp fiction.


The earliest narrative films were rarely more than three to five minutes long, gradually extending to approximately twenty minutes by 1910, and then increasing steadily to a standard feature length of ninety to one hundred twenty minutes by the end of the silent era. Partly to avoid copyright payments and partly to exploit audience familiarity with already existing subject matter at a time when a coherent story could rarely be told on film without the use of copious intertitles or the services of a lecturer within the auditorium to explain the plot, the first adaptations were almost invariably taken from classic authors such as Shakespeare, Dickens, George Eliot (1819–1880), and Thomas Hardy (1840–1928) in Britain, and, on the Continent, Émile Zola (1840–1902), Johann Wolfgang von Goethe (1749–1832), Leo Tolstoy (1828–1910), Alexander Pushkin (1799–1837), and others. The sheer length of most of these works, however, prohibited any attempt at completeness, and standard practice was to choose well-known extracts or scenes that were relatively self-sufficient, such as the "Dotheboys School" scenes from Nicholas Nickleby or the shipwreck scene from The Tempest. As films gradually increased in length, valiant attempts were made to squeeze the whole plot of a novel or film into a running time of around twenty minutes. Popular titles adapted in this early period included Uncle Tom's Cabin (1903), Frankenstein (1910, and much filmed since, though never, despite such titles as Mary Shelley's Frankenstein [1994], with much authenticity), Robinson Crusoe (1913), Faust (1915), and Don Quixote (1915).

Technically, most of these early films were static—filmed from a fixed camera position, usually in long shot, and presenting action in tableau-like form. By the 1910s, however, cinematic technique had become much more sophisticated, with extensive camera movement, fuller use of screen space and camera angle and distance, a more naturalistic acting style, and creative editing that enhanced understanding of plot and character rather than simply moving the action from one setting to another. It became possible to tell stories on the screen with more completeness and complexity, though the desire to give the young medium cultural respectability led to continued reliance on Shakespeare and Dickens in particular. Soon, however, more recent "best-selling" works began to appear on the screen, such as Mrs. Henry Wood's (1814–1887) melodrama East Lynne, filmed as the first British six-reeler (sixty to seventy minutes) in 1913, and, more controversially, D. W. Griffith's (1875–1948) adaptation of Thomas Dixon's (1864–1946) The Clansman, filmed as The Birth of a Nation, one of the longest American features to date, in 1915. By the 1920s, such works predominated, with adaptations of now largely forgotten writers such as "Ouida" (1839–1908), Marie Corelli (1855–1924), Sir Hall Caine (1853–1931), E. Phillips Oppenheim (1866–1946), and the "sensational" novels of such writers as Michael Arlen (1895–1956), whose The Green Hat was filmed as A Woman of Affairs in 1928, starring Greta Garbo (1905–1990); while the endlessly prolific Edgar Wallace (1875–1932) may well hold the record for being the most frequently filmed English-speaking author ever.

In Europe the epics of the Polish novelist Henryk Sienkiewicz (1846–1916), such as Quo Vadis? (filmed in 1912), helped to provide material for the influential Italian historical dramas, and the novels of Selma Lagerlöf (1858–1940) were crucial sources for the great films of Victor Sjöström (1879–1960) and Mauritz Stiller (1883–1928) in Sweden, particularly the former's Körkarlen (The Phantom Carriage, 1921) and the latter's Gösta Berlings saga (1924). In France Jean Renoir's (1894–1979) Nana (1926), Jacques Feyder's (1885–1948) Thérèse Raquin (1928) and Marcel L'Herbier's (1888–1979) L'argent (Money, 1929) were all based on works by the still controversial Zola. L'Herbier also filmed Luigi Pirandello's (1867–1936) Feu Mattias Pascal (The Late Mathias Pascal, 1925) and Feyder adapted both the best-seller L'atlantide (Lost Atlantis, 1920) by Pierre Benoît (1886–1962) and Crainquebille (Bill, 1922) by the then prestigious Anatole France (1844–1924). What is probably the greatest French film of the 1920s, however, was a different sort of adaptation: every word of Carl Theodor Dreyer's (1889–1968) La Passion de Jeanne d'Arc (The Passion of Joan of Arc, 1928) was scrupulously based on the original transcripts of Joan's trial, and the austerity of the filmmaking style exactly matched the sparseness of the dialogue.


While few people today would care whether The Green Hat was in any way betrayed by its transformation into the Garbo vehicle A Woman of Affairs, the situation is very different with an acknowledged literary classic, where readers tend to have fixed, and widely differing, views of the appearance of the characters or setting—not to mention the meaning or interpretation of the work as a whole—and naturally wish to see these perceptions respected on the screen.

There are many other problems too. Even a relatively short novel cannot be filmed word for word within the confines of the two- to three-hour limit of the average film (though Erich von Stroheim [1885–1957] claimed to have done so with his original cut of Greed [1924] from Frank Norris's [1870–1902] novel McTeague). Selection, omission, and condensation of some kind is inevitable. This normally involves suppression of minor characters and subplots, though these may be among the aspects of the book most cherished by readers. More seriously, although a ten-second shot in a film can often replace pages of description of character, landscape, or a house interior, it is rarely possible for a film to convey the detailed analysis of character psychology or motivation crucial to much of the finest fiction without resorting to lengthy stretches of dialogue. Dialogue itself is also a problem, for even the most apparently "naturalistic" speech on the printed page can appear stilted on the screen, and the complex sentence structure of a Henry James (1843–1916) or William Faulkner (1897–1962) is almost impossible to reproduce successfully. Point of view is another difficulty, especially with first-person narration in a novel; film, by its very nature, tends to employ shifting viewpoints throughout and seem to be objective and external rather than internal. Few of these obstacles are ultimately insuperable; they involve a thorough rethinking by the scriptwriter and director and a readiness to substitute techniques appropriate to film for those less suited to it—for example, Harold Pinter's (b. 1930) and Karel Reisz's (1926–2002) film The French Lieutenant's Woman (1981) after John Fowles's (1926–2005) novel.

Adaptations of short stories, on the other hand, present almost exactly opposite problems, for even a long (twenty- to thirty-page) story has to be expanded to fit the minimum ninety minutes of screen time. As a result, incidents barely referred to in the story may be expanded or others invented, new characters may be introduced, plot elements concocted, and brief conversations may be lengthened or new ones created. Though few classic stories can survive this treatment without severe distortion of the original work, some authors have occasionally been better served by adaptations of shorter works than by the treatment of their novels. The Fallen Idol (1948), directed by Carol Reed (1906–1976) from Graham Greene's (1904–1991) story "The Basement Room"; The Rockinghorse Winner (1950), directed by Anthony Pelissier (1912–1988) from the D. H. Lawrence (1885–1930) story; Tomorrow (1972), directed by Joseph Anthony (1912–1993) from the William Faulkner story; and The Innocents (1961), directed by Jack Clayton (1921–1995) from Henry James's "The Turn of the Screw," are all at least the equal of the often more pretentious feature-length films made from the novels of these authors.

The work of almost every classic English novelist from Daniel Defoe (1660–1731) onward has been filmed at least once, and the same is true in America from James Fenimore Cooper's (1789–1851) The Last of the Mohicans and the stories of Edgar Allan Poe (1809–1849) onward. In France, Stendhal (1783–1842), Honoré de Balzac (1799–1850), Gustave Flaubert (1821–1880), Victor Hugo (1802–1885), and Zola have been constant favorites. Possibly the finest adaptations of French literature have been from the novels of Georges Bernanos (1888–1948), where Robert Bresson (1901–1999), in Journal d'un curé de campagne (Diary of a Country Priest, 1950) and Mouchette (1967), has provided the perfect equivalent in cinematic terms of the mood, theme, and characterization of the originals, while Maurice Pialat's Sous le soleil de Satan (Under Satan's Sun, 1987) delivers great emotional power. The inherently "cinematic" novels of Georges Simenon (1903–1989) have been frequently filmed, in France and elsewhere, with Les fiançailles de M. Hire directed strikingly well by both Julien Duvivier (1896–1967) in Panique (Panic, 1946) and Patrice Leconte (b. 1947) in Monsieur Hire (1989).

Adaptations of classic Russian literature during the Soviet period tended to be hampered by excessive respect for the originals, though Sergei Bondarchuk's (1920–1994) version of Tolstoy's Vonya i mir (War and Peace, 1968)—like King Vidor's (1894–1982) American production in 1956—provided a certain degree of visual interest. Anna Karenina has also been frequently filmed, usually in simplified form, and used as a Garbo vehicle in 1935. Iosif Kheifit's film of Anton Chekhov's (1860–1904) story "The Lady with the Little Dog" (Dama s sobachkoy, 1960) was well received abroad. Most films of Fyodor Dostoyevsky's (1821–1881) fiction—including even Akira Kurosawa's (1910–1998) Hakuchi (The Idiot, 1951)—have been unmemorable, with the striking exception of Bresson's Quatre nuits d'un rêveur (Four Nights of a Dreamer, 1971), from the story "White Nights" (also filmed by Luchino Visconti [1906–1976] as Le notti bianche in 1957; restored version 1997) and, especially, Une femme douce (1968) from the story "A Gentle Creature," both of which, despite updating the settings, are typically near-perfect re-creations of mood, character, and theme, while being thoroughly "Bressonian" throughout.

From German literature, R. W. Fassbinder's (1946–1982) 1974 film of Theodor Fontane's Effi Briest surprised many with the director's unusually sober and restrained visual style and sympathetic treatment of the heroine's fate, both aspects re-creating the book with considerable effectiveness. And Eric Rohmer's (b. 1920) version of Heinrich von Kleist's novella "Die Marquise von O …" (The Marquise of O, 1970) transferred successfully to film the author's ironic and tongue-in-cheek presentation of the heroine's bizarre predicament in finding herself pregnant with no memory of any sexual encounter. Thomas Mann's (1875–1955) novella "Death in Venice," however, was controversially filmed by Visconti in 1971 (Morte a Venezia). Some critics gushed over the visual lushness of the setting and Dirk Bogarde's (1921–1999) fine performance, while others objected to the liberties taken with the central character and the awkward attempts at conveying the aesthetic and philosophical themes of the story. By contrast, Visconti's earlier film of Giuseppe di Lampedusa's (1896–1957) Il gattopardo (The Leopard, 1963), especially in its recent fully restored version in 1996, is a masterpiece both of filmmaking and adaptation, brilliantly re-creating both the period setting and the moral and political dilemmas faced by the main character. Other major Italian successes are Bernardo Bertolucci's (b. 1941) Strategia del rango (The Spider's Stratagem, 1970), from a story by Jorge Luis Borges (1899–1986), and Il conformista (The Conformist, 1970) from Alberto Moravia's (1907–1990) novel, with both films expressing their director's personal vision.

The first Japanese film to achieve international success, Akira Kurosawa's Rashomon (1950), was based on two stories by Ryunosuke Akutagawa (1892–1927). The classic novels of Jun'ichiro Tanizaki (1886–1965) and Yasunari Kawabata (1899–1972) have provided source material for several films by Kon Ichikawa (b. 1915) and Mikio Naruse (1905–1969) respectively, while Hiroshi Teshigahara (1927–2001) has specialized in adapting the idiosyncratic fiction of Ko®ô Abe (1924–1993), with Suna no onna (Woman in the Dunes, 1964) becoming an international art house favorite.

Charles Dickens has been the most frequently filmed of classical English novelists, followed, especially in the 1990s, by Jane Austen, Henry James, Thomas Hardy, and E. M. Forster (1879–1970). Each of Austen's six novels has been filmed, either for the cinema or for television, with the most acclaimed versions being Sense and Sensibility (Ang Lee, 1995), Persuasion (Roger Michell, 1995), and the television Pride and Prejudice (also 1995), which compares favorably with the still popular 1940 version starring Greer Garson (1908–1996) and Laurence Olivier (1907–1989). The updating of Emma as Clueless (1995) retains many of Austen's themes but sets them in the context of a contemporary American high school.

The adaptations of E. M. Forster and Henry James by the team of Ismail Merchant (1936–2005) and James Ivory (b. 1928) have often been dismissed as "Masterpiece Theatre" material for their emphasis on accuracy of costume and setting and their close adherence to the details of characterization and plot at the expense of deeper thematic concerns, thus providing merely an agreeable illustration of the text rather than an interpretation of it. Perhaps in reaction to the Merchant-Ivory approach, several recent versions of James's works have attempted to modernize and make explicit what is left unsaid, and to the reader's imagination, in the originals, most obviously in The Portrait of a Lady (Jane Campion, 1996) and The Wings of the Dove (Iain Softley, 1997); Mansfield Park (Patricia Rozema, 1999) has been accused of imposing an overtly political meaning on a nonpolitical text, and Vanity Fair (Mira Nair, 2004) turns William Makepeace Thackeray's (1811–1863) manipulative and possibly murderous Becky Sharp into a feminist heroine.

Other English classic authors frequently filmed include Emily (1818–1848) and Charlotte Brontë (1816–1855), with William Wyler's (1902–1981) 1939 version of Wuthering Heights, despite dealing with only half of the book, being still the most powerful and atmospheric treatment, and the 1944 Jane Eyre maintaining its superiority to most recent versions. Thomas Hardy has been well served by Far from the Madding Crowd (John Schlesinger, 1967), Tess (Roman Polanski, 1979), and Jude (Michael Winterbottom, 1996). The exquisitely beautiful Barry Lyndon (Stanley Kubrick, 1975) catches perfectly the sense of waste and decay beneath the glittering surface of the worlds of high society and war central to Thackeray's novel. From the eighteenth century, Henry Fielding's (1707–1754) Tom Jones was filmed as a high-spirited romp by Tony Richardson (1928–1991) in 1963, an approach that captures one aspect of the novel but far from all of it, and Daniel Defoe's Robinson Crusoe has been filmed often, most surprisingly—and effectively—by Luis Buñuel (1900–1983) (Las adventuas de Robinson Crusoe, 1954).

Among the "moderns" Graham Greene heads the list, though his novels have rarely been filmed with much success apart from the 1947 Brighton Rock, and it is strange that so inherently cinematic a novelist should have been so poorly served on film. Of the two versions of The Quiet American (1958 and 2002) and The End of the Affair (1955 and 2004), the more recent of each title has been the more successful, but Greene still awaits his ideal adaptor. Joseph Conrad (1857–1924) and D. H. Lawrence, whose works have frequently been adapted to film, have rarely been re-created successfully. Alfred Hitchcock's (1899–1980) film of Secret Agent, titled Sabotage (1936), is more Hitchcock than Conrad, and Christopher Hampton's 1996 version is more respectful than inspired. Much the same is true of probably the best of the Lawrence adaptations, the 1960 Sons and Lovers, while Ken Russell's (b. 1927) Women in Love (1969) is better suited to fans of the director than of the author. The fiction of a supposedly lesser author, W. Somerset Maugham (1874–1965), has fared better,

b. Nevada, Missouri, 5 August 1906, d. Newport, Rhode Island, 28 August 1987

John Huston, the son of the actor Walter Huston, was a boxer, actor, and journalist before becoming a scriptwriter and then writer/director. Almost all his films were based on literary sources, ranging from established literary greats such as James Joyce, Herman Melville, Rudyard Kipling, and Dashiell Hammett to other largely forgotten authors. His directorial career began with a masterpiece of both filmmaking and adaptation, The Maltese Falcon (1941), and it ended with another, The Dead in 1987.

Because he drew on such a wide variety of sources, it is difficult to identify "auteurist" elements in Huston's work. Critics generally pick out such themes as group endeavours and quests (often criminal) that fail as a result of moral flaws—particularly greed and self-interest—among the participants. This view applies to some of his best work, such as The Maltese Falcon, The Treasure of the Sierra Madre (1948), The Asphalt Jungle (1950), and The Man Who Would Be King (1975), though not to the majority of his other films. As someone given considerable freedom to choose his own projects, Huston seems to have rather randomly decided on works that appealed to him personally (as with the boxing theme of Fat City, 1972) or gave him the chance to travel to exotic foreign locations (The African Queen, 1951, and The Roots of Heaven, 1958).

Huston's "invisible" camera style is generally subordinated to presentation of character and plot, although lighting, camera angles, editing, close-ups, gesture, movement, and the use of space are never mechanical and always contribute to understanding and responding to the film's meaning. In his color films especially, however, Huston often conducted daring and controversial experiments, as in the attempt in Moulin Rouge (1952) to re-create the ambience of Henri de Toulouse-Lautrec's paintings. Reflections in a Golden Eye (1967) drained every color except red from the image to produce an overall golden glow that was promptly restored to full color by an outraged studio. One of his finest films, Wise Blood (1979), uses distorted camera angles and unnatural color effects to create the bizarre world of Flannery O'Connor's novel and its half-crazed main character.

Huston was also prepared to alter plot and characterization where necessary. The characters played by Humphrey Bogart and Katharine Hepburn in The African Queen are markedly different from those of the novel, and the book's ending is altered to make the quest succeed (for once). In The Asphalt Jungle, Dix Handley, the "hooligan" played by Sterling Hayden, is presented with far more sympathy than in W. R. Burnett's novel, and the closing scene in which Dix dies in a field surrounded by his beloved horses is far more moving than Burnett's more prosaic ending and remains one of the most memorable images in all of Huston's work


The Maltese Falcon (1941), The Treasure of the Sierra Madre (1948), Key Largo (1948), The Asphalt Jungle (1950), The Red Badge of Courage (1951), The African Queen (1951), Moby Dick (1956), The Unforgiven (1960), The Night of the Iguana (1964), Reflections in a Golden Eye (1967), Fat City (1972), The Man Who Would Be King (1975), Wise Blood (1979), Under the Volcano (1984), Prizzi's Honor (1985), The Dead (1987)


Brill, Lesley. John Huston's Filmmaking. Cambridge, UK, and New York: Cambridge University Press, 1997.

Grobel, Laurence. The Hustons. New York: Scribners, 1989.

Hammen, Scott. John Huston. Boston: Twayne, 1985.

Kaminsky, Stuart. John Huston: Maker of Magic. Boston: Houghton Mifflin, 1978.

Long, Robert Emmet, ed. John Huston: Interviews. Jackson: University Press of Mississippi, 2001.

Graham Petrie

in such films as The Letter (1940) and Of Human Bondage (1934).

Classic American fiction has been less fortunate, on the whole. Victor Sjöström's 1926 film of Nathaniel Hawthorne's (1804–1864) The Scarlet Letter, starring a luminous Lillian Gish, is still by far the best version of that book. Clarence Brown's (1890–1987) silent version of Cooper's The Last of the Mohicans (1920) is much superior to any later version, while films based on Mark Twain's (1835–1910) work, such as The Adventures of Tom Sawyer (1938, 1968 [TV]) or The Adventures of Hucklebrry Finn (1939, 1960, 1985 [TV]) have generally been intended for children. John Huston (1906–1987) made a brave but doomed attempt at Herman Melville's (1819–1891) Moby Dick in 1956; Billy Budd (1962), based on a much shorter work, directed by Peter Ustinov (1921–2004) and starring an appropriately angelic Terence Stamp (b. 1938), was more successful. The stories of Edgar Allan Poe have provided the basis for a whole series of films, notably for American International Pictures in the 1960s and 1970s, with few having much connection with the stories beyond the title, yet often, as with The Masque of the Red Death (1964) providing stylish and sophisticated entertainment. Edith Wharton's (1862–1937) The Age of Innocence was, somewhat unexpectedly, turned into a film in 1993 that was both very close to its source and yet paralleled Martin Scorsese's (b. 1942) more typical world of low-life gangsters with their own hierarchies, rituals, and penalties for refusing to conform.

The major figures of twentieth-century American fiction have also been unevenly treated. Faulkner's novels have generally proved remarkably resistant to adaptation, while Clarence Brown's Intruder in the Dust (1949), from one of the author's less complex works, was an effectively straightforward treatment. Films based on Ernest Hemingway's (1899–1961) fiction have fared best when they depart drastically from the original, as with Howard Hawks's (1896–1977) To Have and Have Not (1944) or Robert Siodmak's (1900–1973) expansion of the story The Killers (1946). John Steinbeck's (1902–1968) The Grapes of Wrath provided the basis for John Ford's classic but not particularly faithful film in 1940, and East of Eden (1955) is memorable mostly for the performance of James Dean (1931–1955) under the somewhat over-heated direction of Elia Kazan (1909–2003), who also directed (more sedately) F. Scott Fitzgerald's (1896–1940) unfinished The Last Tycoon (1976). Neither the 1949 nor the 1974 version of The Great Gatsby is considered to be truly successful, despite the meticulous attention to period detail in the latter. The best films adapted from American literature, in fact, have come from works originally considered marginal or beneath serious literary attention.


Dickens has been by far the most filmed of English novelists, with something like one hundred versions in the silent era alone, and numerous further adaptations for both film and television, continuing to the present day. The earliest films could cope only with well-known incidents or brief character sketches from the books; the sheer length of the major novels has always proved a serious stumbling block. It was natural, then, that the first attempts at full-length treatment would be with shorter works such as A Christmas Carol, A Tale of Two Cities, or Oliver Twist, all filmed several times each before 1920.

Though Dickens has often been called the most cinematic of novelists, his books are far from easy to film satisfactorily. The mixture of realism and symbolism, especially in the later novels, the often larger-than-life or grotesque characters, the first-person narration of some books, the pervasive authorial narrative tone and commentary of others, the sheer scope and variety of characters, incidents and settings, and the insistent social and moral analysis of the later works in particular, all provide formidable barriers that have rarely been totally overcome. All of the thirteen novels have been filmed at one time or another, but the choice has consistently been skewed toward the more realistic, usually early, works, or to those that contain the best-known characters—where the filmmaker is often assisted by the illustrations of George Cruikshank (1792–1878) and "Phiz" (Hablot Knight Browne) (1815–1882), which accompanied the original publications. The complex, densely structured, darker books like Bleak House, Little Dorrit, and Our Mutual Friend have generally met with far less favor.

Though few, if any, of the film adaptations have coped with all the challenges presented by the books, there have been several at least partial successes. David Copperfield, A Tale of Two Cities, A Christmas Carol, Oliver Twist, and Great Expectations have been the most frequently filmed, with, in almost every case, the focus being fixed on character and plot rather than the social criticism that made Dickens such an important figure in his time. The most notable of these include the MGM David Copperfield of 1935, sensitively directed by George Cukor (1899–1983) and with inspired casting that included W. C. Fields (1880–1946) as Micawber, and the same studio's A Tale of Two Cities (also 1935), with a memorable performance by Ronald Colman (1891–1958) as Sydney Carton. These two films still stand as the best adaptations of these books. David Lean's (1908–1991) Great Expectations (1946) and Oliver Twist (1948) are generally considered the classic treatments of these works and the definitive A Christmas Carol is widely acknowledged to be the 1951 Scrooge, starring Alastair Sim (1900–1976). Though Lean's Great Expectations is often considered the finest of Dickens adaptations, it can be argued that his version of Oliver Twist succeeds better in capturing the many dimensions of Dickens's work—the realistic, the grotesque, the comical, the social comment, the sentimental, the symbolic, the fascination with violence—presented in imagery that creates London both as a real city and a symbolic underworld. It does all this much more successfully than Polanski's disappointing treatment (2005). Other interesting versions of less frequently filmed works include The Mystery of Edwin Drood (Stuart Walker, 1935), Nicholas Nickleby (Alberto Cavalcanti, 1947), and the ambitious but flawed two-part Little Dorrit (Christine Edzard, 1988). The well-cast and intelligently reworked Nicholas Nickleby (Douglas McGrath, 2002) unfortunately met with scant interest at the box office. In recent years the most impressive adaptations have come from British television, where the serial format of three to four hours or more can allow a fuller and more leisurely treatment of the texts. Some of the best of these have been Granada Television's Hard Times (1977) and the BBC's Bleak House (1985), Martin Chuzzlewit (1994), and Our Mutual Friend (1998)—all of them books largely neglected by the cinema.

Although all the films mentioned are set in the Victorian period, there have been some attempts at updating them. Rich Man's Folly (1931), a truncated and unsatisfactory version of Dombey and Son, is set at the time of filming, as is a misbegotten Great Expectations (Alfonso Cuarón, 1998), which succeeds in getting almost everything about the novel wrong. By far the best updating is the Portuguese director Joâo Botelho's (b. 1949) Tempos difíceis (Hard Times, 1989), where Dickens's assault on the capitalist mentality remains as relevant today as it was during his lifetime. And, although most of the films based on Dickens's works have come from the English-speaking world, there have also been German, French, Italian, Danish, Russian, and Hungarian treatments, mostly in the silent period.


American cinema is largely a genre cinema. Melodramas, westerns, crime and gangster films, science fiction films, historical and biblical epics, comedies, war films, and musicals have formed the staple of its offerings from the very beginning. A surprising number of these are based on written sources, but because most of these are not canonical in the way that the works of Dickens or Austen are, this goes largely unnoticed and scant attention is paid to whether they have been faithfully adapted or not. As almost all of these genres focus on action, movement, setting (urban or rural), and atmosphere, and generally offer little scope for complexity of character, elaborately phrased dialogue, or intense psychological analysis, they are eminently suited for film.

The inherently "filmic" genre of the western is far more dependent on written sources than is generally realized, ranging from some of the few acknowledged literary classics such as Jack Schaefer's (1907–1991) Shane, filmed by George Stevens (1904–1985) in 1953, to the more ephemeral magazine stories and pulp novels on which films like High Noon (1952) and Stagecoach (1939) were based. In these and similar cases, little more than a basic plot and some aspects of character and setting are generally all that is taken over from source to film.

Crime and gangster films, including films noirs, are also heavily indebted to literary sources, many of them now gaining belated critical respect. Here, too, a considerable laxity in transformation from book to film has been widespread, even with major writers such as Raymond Chandler (1888–1959) and Dashiell Hammett (1894–1961), where only The Maltese Falcon (1941) has survived intact in its adapted form. Less "reputable" writers such as James M. Cain (1892–1977), Jim Thompson (1906–1977), Cornell Woolrich (1903–1968), and David Goodis (1917–1967) have nevertheless provided the basis for some of the finest of American (and also French) films, once again in the form of loose or free rather than strict adaptations. Cain's Double Indemnity, The Postman Always Rings Twice (filmed at least four times to date), and Mildred Pierce were turned into 1940s classics, and a sudden vogue for Thompson produced several adaptations in the 1980s and 1990s, the most successful probably being Coup de Torchon (Clean Up, Bertrand Tavernier, 1981), based on Pop. 1280, which, despite being set in French colonial Africa rather than the American South, brilliantly captures the sleaze, cynicism, and nihilism of the novel. Woolrich, under both that name and William Irish, wrote the original story that Hitchcock filmed, much altered and expanded, as Rear Window (1954), and also the novels on which Hitchcock's admirer François Truffaut (1932–1984) based La marié était en noir (The Bride Wore Black, 1968) and The Mississippi Mermaid (1969), as well as providing the source for such films noirs as Phantom Lady (1944). Truffaut also filmed, with considerable fidelity, Goodis's despairing Down There as Tirez sur le pianiste (Shoot the Pianist, 1960).

The Sherlock Holmes stories of Sir Arthur Conan Doyle (1859–1930) and his novel The Hound of the Baskervilles have been endlessly reworked (or, in some cases, invented) for both film and television, with critical debate centering mainly on who has been the "best" or most "authentic" Holmes or Watson; a similar fate has met Ian Fleming's (1908–1964) James Bond. And a rather neglected figure in crime fiction, W. R. Burnett (1899–1982), provided the original stories on which such classics as Little Caesar (1931), High Sierra (1941), and The Asphalt Jungle (1950) were based.

b. Chicago, Illinois, 23 July 1888, d. La Jolla, California, 26 March 1959

Educated in England, Raymond Chandler worked as an accountant and in a bank on returning to America before turning to writing pulp fiction in the 1930s. The success of his first novel, The Big Sleep (1939), brought him an invitation to Hollywood. His involvement with film had two aspects: as screenwriter and as author of six novels adapted for the screen, some of them more than once. After a rewarding experience collaborating with Billy Wilder on the script of Double Indemnity (1944), Chandler became increasingly disillusioned with Hollywood and attacked it as a soul-destroying environment in articles written for Atlantic Monthly. Apart from receiving cowriting credit on two minor films in 1944 and 1945, his only further completed work for the screen was an original script for The Blue Dahlia (1946). He received only cowriter credit on Alfred Hitchcock's Strangers on a Train (1951) after disagreements with the director.

The first two film versions of his novels, The Falcon Takes Over (1942), loosely based on Farewell, My Lovely, and Time to Kill (1942), based on The High Window, retained only aspects of the plots and created a Philip Marlowe character very different from Chandler's original. A more serious attempt at adapting Chandler's work came in Murder, My Sweet (1944), again from Farewell, My Lovely, with Marlowe played by Dick Powell. This was followed by what is considered to be the finest Chandler adaptation, The Big Sleep (1946), directed by Howard Hawks, with Humphrey Bogart as the definitive Marlowe, even though he played the role only once. The Lady in the Lake (1947) made a largely unsuccessful attempt to use the camera as first-person narrator, with Marlowe seen only in mirrors until the very end of the film. The Brasher Doubloon (1947), a weak adaptation of The High Window, starred George Montgomery as an unconvincing Marlowe.

Twenty years passed before further adaptations were made, creating problems with attempts to re-create the very specific 1940s settings, themes, and ethos of the novels. Marlowe (1969), based on The Little Sister and starring James Garner, updated the story to the 1960s and presented the hero as a figure of integrity who was out of step with the times. Robert Altman's The Long Goodbye (1973) went even further by presenting Elliot Gould as a bewildered and largely ineffectual figure in 1970s Los Angeles—and treated as a figure of fun by most of the other characters. Although the film was disliked by many Chandler admirers, it remains a brilliant piece of filmmaking. The two most recent versions both starred an ageing Robert Mitchum. Farewell, My Lovely (1975) took great pains to re-create the settings and atmosphere of the book, and a Big Sleep (1978), directed by Michael Winner and set bizarrely in contemporary London, suffered fatally by comparison with Hawks's film.


Double Indemnity (1944), Murder, My Sweet (1944), The Big Sleep (1946), The Blue Dahlia (1946), The Lady in the Lake (1947), Strangers on a Train (1951), The Long Goodbye (1973), Farewell, My Lovely (1975)


Clark, Al. Raymond Chandler in Hollywood. London and New York: Proteus, 1982.

Gardiner, Dorothy, and Kathrine Sorley Walker, eds. Raymond Chandler Speaking. Boston: Houghton Mifflin, 1977.

Luhr, William. Raymond Chandler and Film. New York: Frederick Ungar, 1982.

Pendo, Stephen. Raymond Chandler on Screen: His Novels into Film. Metuchen, NJ: Scarecrow, 1976.

Graham Petrie


Film historians have noted the close links between theatrical melodrama of the late nineteenth century and the techniques and narrative structure of early film—in content and elaborate lighting and stage effects. The obvious similarities between a play and a film—in overall length, use of sets, the apparent realism of character and dialogue—have obscured the very real differences. Stage dialogue can sound artificial and tedious when transferred directly to the more naturalistic medium of film, and, as with fiction, a successful adaptation has to be thoroughly rethought in terms of the new, primarily visual, medium of cinema. While the faults of mechanically adapted "filmed theater" are usually obvious, there

is equal danger in attempts to "open out" a play by transferring interior scenes into exotic outdoor locations and hoping that will somehow make the work more cinematic. Some sort of balance between stage and film effects is therefore essential. Sidney Lumet's (b. 1924) filming of Eugene O'Neill's (1888–1953) Long Day's Journey into Night (1962) achieves its claustrophobic effect by respecting the spatial limitations of the stage while transforming it through skillful use of camera movement and lighting, and by varying screen space and distance for dramatic effect.

Shakespeare has been by far the most adapted playwright worldwide, even in the silent period, when extracts and condensed versions of his plays proliferated in most European countries as well as in Britain and the United States. The coming of sound brought the inevitable problem of how to make poetic dialogue convincing in the more naturalistic medium of film. It is often argued that the finest of all Shakespeare films is Kurosawa's 1957 Kumonosu jô (Throne of Blood), which is based on Macbeth. It retains almost nothing of the dialogue, even in Japanese, while majestically transforming theme, emotion, and imagery into purely visual terms, with Macbeth constantly surrounded by images of fog, nets, and labyrinths. Though Grigori Kozintsev's (1905–1973) Gamlet (Hamlet, 1964) and Korol Lir (King Lear, 1970) use Boris Pasternak's (1890–1960) translation of the plays, the non-Russian–speaking viewer, forced to rely on subtitles, can perhaps appreciate better the stark black-and-white imagery of the films.

The most admired English-language versions usually attempt a compromise between stylization and naturalism, both in speech and action; for example, Laurence Olivier used the confined space of the castle set in Hamlet (1948) and allowed the camera full rein in the battle scenes of Henry V (1944). Polanski's Macbeth (1971) accentuates the physical violence inherent in the play, and Orson Welles (1915–1985) brings his own superb visual sense to his Othello (1952) and Campanadas a medianoche (Chimes at Midnight, 1967, based on the Henry IV plays) without neglecting the spoken word. Examples of more radical transformations are the updating of Romeo and Juliet by Baz Luhrmann (1996) and the intensely personal re-creations of The Tempest (1979) by Derek Jarman (1942–1994) and Peter Greenaway (b. 1942) (as Prospero's Books, 1990). Kenneth Branagh (b. 1960), in seemingly open competition with Olivier, has filmed an uncut Hamlet (1996) and an impressive Henry V (1989), among others.

The most often filmed English dramatists after Shakespeare have been George Bernard Shaw (1856–1950), Noel Coward (1899–1973), Terence Rattigan (1911–1977), and Oscar Wilde (1856–1900). In most cases the results have been respectful and moderately faithful rather than inspired (though the 1928 film of Coward's The Vortex and the 1933 Design for Living had to be drastically altered to escape the censors). Anthony Asquith's (1902–1968) 1952 film of The Importance of Being Earnest still far surpasses later versions of Wilde, both as a film and as an adaptation, and both versions of Rattigan's The Browning Version (1951, 1994) and The Winslow Boy (1948, 1999) remain popular.

Eugene O'Neill, Tennessee Williams (1911–1983), Arthur Miller (1915–2005), Clifford Odets (1906–1963), and Lillian Hellman (1906–1984) are among the most frequently adapted American playwrights, though, with Williams in particular, contentious subject matter has often forced major alterations between stage and screen. A Streetcar Named Desire, directed by Elia Kazan in 1951, remains the classic transformation of his work. Apart from the version of Long Day's Journey into Night, the best O'Neill adaptation has been John Frankenheimer's (1930–2002) The Iceman Cometh (1975). Hellman's The Little Foxes (1941) became a classic film through William Wyler, but Clash by Night (1952) and The Big Knife (1955) are largely rewritten versions of Odets. Perhaps the most interesting film based on Arthur Miller's work is Sorcières de Salem (TheWitches of Salem, 1957), from The Crucible, with a script by Jean-Paul Sartre (1905–1980).

In Europe, Henrik Ibsen (1828–1906), August Strindberg (1849–1912), and Anton Chekov (1860–1904) have often been adapted. The 1951 Fröken Julie (Miss Julie), directed by Alf Sjöberg (1903–1980), is still the best Strindberg, but few of the English-language films of Ibsen and Chekov have been particularly successful. Jean Renoir (Les bas-fonds, 1936) and Akira Kurosawa (Donzoko, 1957) made very different but equally fascinating films of Maxim Gorky's (1868–1936) The Lower Depths.


Detstvo Gorkogo (The Childhood of Maxim Gorky, 1938), directed by Mark Donskoy (1901–1981), remains one of the finest of film biographies/autobiographies, but most such films are bedevilled by questions of authenticity, for content is more important here than transforming sophisticated literary techniques into film. Does the leading actor really resemble the subject (whose photos or portraits are usually well known)? Is the film factually accurate or truthful (and is this true of its source)? Is it slanted in favor of or against the protagonist? Are there distortions of fact, omissions, invented incidents or encounters? Some film biographies, such as Finding Neverland (2004), admit to not being completely factual, but most do not, and the majority of such films are built up by drawing on a variety of sources, augmented by scenes imagined or created by the scriptwriter. The result, as in Martin Scorsese's Raging Bull (1980), may be superb cinema but should not necessarily be considered a definitive account of the subject's life.

Comic books and comic strips have proved a consistent source of film material, though the various treatments of Batman and Superman, for example, usually consist of rewritten works based on a variety of incidents taken from the original rather than an adaptation of one particular story. Many popular television series have been turned into films, such as The Addams Family (1991) or The Brady Bunch (1995), on much the same principle of selection, and the recent vogue for graphic novels has also spilled over into film, as with Ghost World (2001) from the original by Daniel Clowes (b. 1961).

Films for children tend to be either live action, as in the several versions of Little Women (1933, 1949, 1994) and The Secret Garden (most recently 1993), or animated, as with the Disney classics Snow White and the Seven Dwarfs (1937) and Bambi (1942), though more recent films from that studio are too often saccharine distortions of what were quite tough-minded originals. The digital animation of The Polar Express (2004) recreates the visual world of the book very convincingly. Opera on film tends to be similar to "canned theater" (Miss Julie), directed by Alf Sjo with a few exceptions, such as Joseph Losey's (1909–1984) Don Giovanni (1979) or Francesco Rosi's (b. 1922) Carmen (1984), which were well reimagined for film. And longer poems such as Henry Wadsworth Longfellow's (1807–1882) Hiawatha (1952) or Alfred Lord Tennyson's (1809–1892) The Charge of the Light Brigade and Geoffrey Chaucer's (1340–1400) The Canterbury Tales have become (very loosely) the basis for feature-length films. Overall, then, almost anything written, or even drawn, can be transformed into a film, either faithfully or altered almost out of recognition, with success depending as much on the skill and intelligence of the filmmaker as the often uneven quality of the original material.

SEE ALSO Biography;Comics and Comic Books;Screenwriting;Theater


Ball, Robert Hamilton. Shakespeare on Silent Film: A Strange, Eventful History. New York: Theater Arts Books, 1968.

Bazin, André. What Is Cinema? 2 volumes. Translated by Hugh Gray. Berkeley: University of California Press, 1967.

Bluestone, George. Novels into Film. Berkeley: University of California Press, 1973 [1957].

Chatman, Seymour. Coming to Terms: The Rhetoric of Narrative in Fiction and Film. Ithaca, NY: Cornell University Press, 1990.

Enser, A. G. S. Filmed Books and Plays: A List of Books and Plays from which Films Have Been Made, 1928–86. Hampshire, UK: Gower Publishing, 1987.

Giddings, Robert, Keith Selby, and Chris Wensley. Screening the Novel: The Theory and Practice of Literary Dramatization. New York: St. Martin's Press, 1990.

Jorgens, Jack J. Shakespeare on Film. Bloomington: Indiana University Press, 1977.

Klein, Michael, and Gillian Parker, eds. The English Novel and the Movies. New York: Frederick Ungar, 1981.

Naremore, James, ed. Film Adaptation. New Brunswick, NJ: Rutgers University Press, 2000.

Peary, Gerald, and Roger Shatzkin, eds. The Classic American Novel and the Movies. New York: Frederick Ungar, 1977.

——, eds. The Modern American Novel and the Movies. New York: Frederick Ungar, 1978.

Sinyard, Neil. Filming Literature: The Art of Screen Adaptation. New York: St. Martin's Press, 1986.

Smith, Grahame. Dickens and the Dream of Cinema. New York: Manchester University Press, 2003.

Smith, Murray. Engaging Characters: Fiction, Emotion, and the Cinema. New York: Oxford University Press, 1995.

Wheeler, David, ed. No, But I Saw the Movie: The Best Short Stories Ever Made into Film. New York: Penguin Books, 1989.

Graham Petrie

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An adaptation to climate change is any adjustment in a natural or human system that decreases the harm caused by climate change or takes advantage of some opportunity it offers. Both natural and human systems may adapt to actually occurring climate change; humans are unique in being able to forecast changes and attempt to adapt to them in a planned way.

There are three types of adaptation, namely anticipatory, autonomous, and planned. Anticipatory adaptation takes place before predicted impacts are observed; autonomous or spontaneous adaptation takes place in direct response to ecological or economic impacts of climate change; and planned adaptation is based on deliberate policy decisions. How much adaptation will occur depends on several factors, including how much climate changes, how severe the impacts of that change are, where they are most severe, how adaptable impacted ecosystems and human communities are, and whether effective policies are followed for mitigating and adapting to climate impacts.

Some impacts of climate change are already being felt, and some adaptation by both natural and human systems has already begun. However, adaptation is not always possible. For ecosystems, failure to adapt means the collapse of existing ecologies and their replacement by others, along with species extinctions. For human communities, failure to adapt means partial or severe breakdown of existing ways of life, with consequences ranging from monetary losses to famine and mass death. In general, slower climate change is easier to adapt to than sudden change, and slight change is easier to adapt to than extreme change.

Historical Background and Scientific Foundations

Natural and Historic Adaptation

Over geologic time, Earth's climate has changed regionally and globally many times. Ice ages have spanned millions of years, dotted by glacial and interglacial cycles; the sun has gradually warmed throughout life's history; carbon dioxide (CO2) concentrations were far higher than today's during most of Earth's history; and regional shifts in rainfall patterns and temperature have been common. Adaptation by natural systems—or failure to adapt, on a few occasions leading to mass extinctions involving the majority of living species—has thus always been a feature of life on Earth.

Since the evolution of human beings, regional and local climate changes have continued to occur, though not with the intensity of the most drastic changes of the deep past. Some climate changes, such as the El Niño– Southern Oscillation climate cycle, occur regularly and last for years, and adaptation to them is routine in affected parts of the world. Affected ecosystems and peoples have, historically, sought to adapt to climate changes, often successfully.

Most plants and animals are evolved to thrive in a certain range of climate conditions: too little or too much precipitation or warmth are not tolerable to them. Therefore no species is found everywhere on Earth, but only where climate conditions are favorable and where the species has managed to cross geographical barriers to colonize a given area. For example, the European downy birch, a tree species, requires cold temperatures. It thrives in northern Asia and Europe and is the only tree native to Greenland and Iceland; it is not found in tropical areas, even in high mountainous zones where it might be able to survive if its seeds could reach those locations.

Natural adaptation for plants consists mostly of range shifting. As global warming proceeds, a plant such as the European downy birch will gradually cease to be found in the southernmost parts of its historical range and will seed itself northward to areas that were formerly too cold. Since plants and animals tend to live in adapted communities, with animals depending on particular plants and vice versa, entire biomes (types of habitat) will shift location as climate changes, plants and animals

together. Such shifts may happen quickly, on the scale of a human lifetime: range shifts have already been measured in many parts of the world.

For mountainous species, as warming causes cold climate zones to move to higher altitudes, habitats move upward and shrink. For example, the uppermost altitude for pine mistletoe in Switzerland was 656 ft (200 m) higher in 2004 than it was in 1910. Range shifts can also occur more slowly. In North America, almost all of present-day Canada was covered by ice about 21,000 years ago, and spruce trees were found across what is now the northern and central United States, down into Texas and the Southwest. Over the next 15,000 years, spruce forests migrated northward as the ice retreated. By 7,000 years ago, spruce was common only in northern areas that had formerly been covered by ice. This pattern persists today, and modern climate change will drive spruce forests even farther north.

Over longer time periods, adaptation may be genetic as well as geographic. However, evolving new climate tolerances is a far slower, chancier process than colonizing new regions. Failure to either migrate or evolve leads to extinction.

Human history shows varying levels of adaptation to climate swings. For example, the world's first civilized empire, the Akkadia, was established between 4300 and 4200 BC in Mesopotamia, that is, the plain between the Tigris and Euphrates rivers, much of which is located in what is Iraq today. After only about 100 years of prosperity, the empire collapsed. Soil cores from the region—cylinders of undisturbed soil extracted vertically from the ground—contain a layer of wind-blown silt that records a 300-year drought that began shortly before the Akkadian collapse. Archaeological evidence shows that the Akkadians tried to adapt to this change by building larger grain storage facilities and irrigation systems, but these adaptations were insufficient.

From paleoclimatic evidence, the Mesopotamian drought has been linked to a cooling event in the North Atlantic in which surface water temperatures declined by 1.8–3.6°F (1–2°C). Modern temperature records show that the year-to-year water supply of the Mesopotamian plain can be cut in half by unusual cooling of the Atlantic. Century-scale regional drought has also been implicated in the collapse of the classic Maya civilization around AD 800, along with overpopulation, deforestation, erosion, and war, and of the Tiwanaku culture in the Bolivian-Peruvian altiplano about AD 1000.


ALTIPLANO: High plateau region in the Andes of Argentina, Bolivia, and Peru in South America, the second-largest high plateau on Earth after Tibet. Sediments in Lake Titicaca, into which the Altiplano drains, record tens of thousands of years of regional climate change.

BIOME: Well-defined terrestrial environment (e.g., desert, tundra, or tropical forest). The complex of living organisms found in an ecological region.

DEFORESTATION: Those practices or processes that result in the change of forested lands to non-forest uses. This is often cited as one of the major causes of the enhanced greenhouse effect for two reasons: 1) the burning or decomposition of the wood releases carbon dioxide; and 2) trees that once removed carbon dioxide from the atmosphere in the process of photosyn-thesis are no longer present and contributing to carbon storage.

EL NIÑO–SOUTHERN OSCILLATION: Global climate cycle that arises from interaction of ocean and atmospheric circulations. Every 2 to 7 years, westward-blowing winds over the Pacific subside, allowing warm water to migrate across the Pacific from west to east. This suppresses normal upwelling of cold, nutrient-rich waters in the eastern Pacific, shrinking fish populations and changing weather patterns around the world.

GEOLOGIC TIME: The period of time extending from the formation of Earth to the present.

GLACIAL CYCLE: Episode in Earth climate history in which temperatures decline and glaciers grow and spread, sometimes covering large parts of the northern and southern hemispheres. The most recent glacial cycle ended about 10,000 years ago.

ICE AGE: Period of glacial advance.

MANGROVE FOREST: Coastal ecosystem type based on mangrove trees standing in shallow ocean water: also termed mangrove swamp. Mangrove forests support shrimp fisheries and are threatened by rising sea levels due to climate change.

PALEOCLIMATE: The climate of a given period of time in the geologic past.

RANGE SHIFT: Movement or shrinkage of the territory occupied by a given species of plant or animal due to climate change. When climate warms, species cease to occupy areas that were at the warm extreme of their ability to adapt and to colonize areas that were at the cool extreme. The result is a shifting range.

RIVER DELTA: Flat area of fine-grained sediments that forms where a river meets a larger, stiller body of water such as the ocean. Rivers carry particles in their turbulent waters that settle out (sink) when the water mixes with quieter water and slows down; these particles build the delta. Deltas are named after the Greek letter delta, which looks like a triangle. Very large deltas are termed megadeltas and are often thickly settled by human beings. Rising sea levels threaten settlements on megadeltas.

The collapse of the urban and military structures of these societies can be viewed as a form of adaptation: in response to increased scarcity brought on by climate change, they reduced their social complexity, abandoned cities, and reorganized their systems of production and supply. In all cases, human populations persisted, though decreased in number. However, such forced, autonomous adaptation is painful and often deadly for individuals caught up in it.

The global warming being experienced today is unique in Earth's history in that it is caused by human beings. Today's concern with anticipatory, planned adaptation to human-caused (anthropogenic) climate change is also a historical first. Not until the late twentieth century did humans understand the causes of climate change and possess the ability to make reasonably reliable predictions of its future course. The scientific study of adaptation to sea-level rise and the other effects of climate change began in the late 1980s and early 1990s, when extensive studies of adaptation strategies began to appear.

Modern adaptation, whether autonomous (reacting to existing changes) or anticipatory, will be based on the impacts of climate change. Hundreds of these potential impacts have been studied and many are being observed already, including rising sea levels, shifts in rainfall, increased heat waves, migrations of biomes (regional ecosystems), bleaching of corals, and many more. Impacts are often outlined by geographic region or by sector.

Freshwater Resources

Projected increases in air temperature, variability of rainfall, and sea level will impact freshwater systems in many parts of the world during the coming centuries. Over 16% of the world's population lives in basins fed by glacial melt or snow-melt flows, which will decline as warming shrinks glaciers, decreases precipitation in some areas, and shortens snow seasons. Rising sea level will extend salty water inland by penetrating groundwater aquifers, which will reduce freshwater availability for communities in coastal zones. Increased precipitation in some areas will increase flooding risk; decreased precipitation in others will increase drought risk.

Adaptations to impacts on freshwater resources can be either supply-side or demand-side. Supply-side adaptations seek to ensure supplies, while demand-side adaptations seek to reduce usage, allowing communities to prosper even when supplies decrease. Some supply-side adaptations include desalination of seawater, increasing storage capacity by building reservoirs, and expanding rainwater storage systems. Some demand-side adaptations include water recycling to increase water-use efficiency, reducing irrigation demands by changing crop mixes, planting calendars, irrigation methods, and providing incentives for water conservation such as metering and pricing.


Increased atmospheric CO2, along with the warming that it causes, will affect most ecosystems in the coming century and for long after. By the end of the twenty-first century, atmospheric CO2 exceeds levels seen for at least 650,000 years. Even apart from warming, CO2 has direct ecosystem effects, speeding the growth of terrestrial plants and making the oceans significantly more acidic. Some 20–30% of plant and animal species that had been specifically examined as of 2005 in an unbiased (random) sample were likely to be at increased extinction risk as the world warms by 3.6–5.4°F (2–3°C) above pre-industrial (pre–1750) levels.

Adaptations to impacts on ecosystems can be both natural and human. Natural adaptations will consist mostly of biome migration. Human responses will involve changes in management of natural resources and wild areas. There are a number of ways that human managers can increase the resilience of ecosystems, though these will likely become less effective or outright useless at higher levels of climate change. First, monitoring changes in climate and ecosystems is essential to allow for effective adjustments in management. Reducing harm to natural systems by other human activities—including pollution, habitat destruction, habitat fragmentation (where development and habitat destruction break up habitats into isolated pieces), and the introduction of invasive alien species—will almost always enhance the ability of ecosystems to adapt to climate change.

Expansionofnational forests andparks andother kinds of reserve systems will reduce ecosystem vulnerability to climate change, especially if reserves are designed to consider long-term migrations of biomes and human settlements. For particular species, managing for connected populations (rather than isolated pockets of population), genetic diversity, and larger populations will increase survivability in response not only to climate change but to other challenges.

Agriculture and Forestry

In some mid-latitude and northerly regions, moderate global warming—that is, warming of 1.8–5.4°F (1–3°C)—is likely to benefit agriculture by increasing yields from pastures and crops. However, even mild warming will decrease yields in low-latitude regions, including the tropics, where most of the world's poor live. Higher levels of warming, which are quite possible, will have negative impacts on agriculture in all regions.

Although United Nations scenarios for possible world development show the number of malnourished persons in the world declining from about 820 million today to 100–380 million by 2080, this range is greater than it would be if global climate change were not occurring. Climate change also serves to shift the regional distribution of hunger, especially making hunger worse in sub-Saharan Africa. The productivity of commercial forestry is forecast to rise slightly to modestly in the short and medium term, partly due to the fertilizing affect of CO2 on young trees.

There are many possible adaptive responses to climate change's impacts on agriculture. These include

altering the varieties or species planted to those better adapted to increased warmth or decreased rainfall, more efficient irrigation techniques, altering the timing and location of crop planting, and diversifying income generation (for example, by raising livestock). In a study of agricultural adaptation to hotter, drier summers in Modena, Italy, it was predicted that with unchanged farming practices sorghum crops would be reduced by 48–58%. With adjusted sorghum varieties and planting times, the impact could potentially be reduced to zero.

Coastal Systems

Rising sea levels and increased storm violence will impact coastal areas around the world. Over a billion people live in coastal areas today, with coastal population growing to over 5 billion by 2080 according to some global-development scenarios. The 300 million people living on large river deltas—close to, at, or even below present-day sea level—will be particularly at risk for impacts, especially flooding. Sea level is predicted to rise by 2 ft (0.6 m) or more by 2100, and to continue rising thereafter.

Over the next several centuries to a millennium, complete melting of the Greenland and West Antarctic Peninsula ice sheets could raise sea levels by 40 ft (12 m), radically altering coastlines. Hurricanes and tropical cyclones will probably increase in intensity, and the effects of these intensified storms will be amplified by higher sea levels. Without improved coastal protection, coastal flooding could increase by a factor of 10 by the 2080s, affecting over 100 million people, mostly in developing countries.

Adaptations to the coastal impacts of climate change take three basic forms: protection, accommodation, or retreat. Protection is practically synonymous with the building of dikes to keep back the sea, as in Holland and New Orleans, Louisiana. Accommodation may include flood-resistant building construction (e.g., buildings on pilings) or floating agricultural systems (now being tested in Holland). Retreat essentially means moving settlements back to higher ground. Astute combination of various forms of these adaptation options could reduce the impacts of sea-level rise by 10 to 100 times in many areas; at the other extreme, as for small islands in the Pacific, no adaptation may be feasible, and “retreat” may have to signify abandonment.

Human Health

Climate change is projected to increase malnutrition; injure child growth and development; increase the number of people suffering death and disease from extreme weather events such as storms, droughts, heat waves, and floods; shift the ranges of some infectious disease vectors; increase the amount of diarrheal disease (which presently causes 5–8 million deaths per year, mostly children); increase cardio-respiratory disease due to ground-level ozone; and decrease the number of deaths from cold in northern regions. Hurricane Katrina (2005) showed that even highly developed countries such as the United States may not be well-prepared for the health and other consequences of extreme weather events.

Adaptations to the impacts of climate change on health will mostly take the form of revised policies and procedures of national and international health organizations (in the United States, for example, the Centers for Disease Control and Prevention [CDC]). Climate-based early warning systems for heat waves and malaria outbreaks have already been implemented in some countries. Seasonal forecasts of events such as drought can allow the timely launching of public education campaigns on the prevention of diarrheal and other infectious diseases.

Impacts and Issues

The types and amounts of adaptations required—or possible—will depend on the amount of climate change that occurs. The more effective mitigation efforts, the less adaptation and failed adaptation will occur. Also, the pattern of impacts and therefore of demands on adaptation will be patchy. Tundra, boreal (northern-type) forests, and mountain and Mediterranean ecosystems will be highly vulnerable, as will mangroves and salt marshes along coasts. Low-lying coasts will be more vulnerable than steep coasts, and water resources will be more endangered in mid-latitude and dry tropical regions due to lessened rainfall and increased evaporation. Agriculture will be at risk in tropical regions. Africa, the Arctic, small islands, and large Asian river deltas are the areas most at risk from climate change.


According to the National Academy of Sciences: “Evidence shows that the climate has sometimes changed abruptly in the past—within a decade—and could do so again. Abrupt changes, such as the Dust Bowl drought of the 1930s that displaced hundreds of thousands of people in the American Great Plains, take place so rapidly that humans and ecosystems have difficulty adapting to them.”

SOURCE:Staudt, Amanda, Nancy Huddleston, and Sandi Rudenstein. Understanding and Responding to Climate Change. National Academy of Sciences, 2006.

As a rule, adaptation costs money. Therefore, wealthier nations would be better able to adapt even if they were facing equally grave impacts from climate change—but they are not. Impacts are likely to be much more severe in the developing countries that are less equipped to adapt to them. For example, by 2100, developing countries stand to lose about 10 times more land area to rising sea levels than developed countries, with four million people likely displaced versus a few hundred thousand, at most, in the developed countries.

Uncertainties about what climate change will occur makes planned adaptation more difficult, though there are many measures (called no-regrets measures) that carry benefits regardless of how much climate change occurs. For example, increased wildlands conservation will reduce species extinctions under any future climate scenario. Evaluation of which planned adaptations will be worth their cost depends sensitively on estimates of the social cost of carbon, that is, the total amount of future economic harm that will be caused by each ton of carbon (or equivalent amount of another greenhouse gas) emitted to the atmosphere. Calculation of this value is notoriously elusive and values-laden. The willingness or ability of governments to respond to threats that act on time-scales much greater than the election cycles or a ruler's personal lifespan also threaten timely and effective mitigation of and adaptation to climate change.

Primary Source Connection

Local ecosystems and the safety of human populations are put at risk as humans modify the environment around them. Local and national governments can minimize these effects by implementing adaptation responses. An adaptation response is a plan that addresses foreseeable environmental issues. Examples of adaptation responses include forest management, flood control, and habitat preservation and restoration. This article examines some of the adaptation responses taken by several European countries.

The European Environment Agency (EEA) is an agency of the European Union devoted to monitoring the environment in Europe and promoting sustainable development. The EEA has 32 member countries and six cooperating countries.


As a region of industrialised nations, Europe has a strong commitment to mitigating climate change by reducing greenhouse gas emissions. However, numerous scientific studies, as summarised in Section 3 of this report, and the considerable losses resulting from extreme weather events over recent years…demonstrated the vulnerability of Europe's natural environment and its society to projected climate change impacts. There is growing recognition that Europe should adapt to such impacts in order to maintain sustainable functioning of ecosystems and wellbeing of its population. Many EEA member countries have started to adjust their overall national climate policy framework to include climate change adaptation as an equally important component as mitigation. A wide range of adaptation responses have been initiated at varying governmental levels and in different sectors….

This section reviews adaptive responses in member countries in relation to natural ecosystems, water resources, coastal and river floods, natural hazards, human health and the business sector.

4.4.1 Maintaining the health of Europe's ecosystems

To address the wide ranging adverse impacts of climate change to Europe's terrestrial ecosystems, a variety of measures have been initiated or are being planned. This is often carried out in the context of nature conservation and sustainable resource management rather than deliberately directed at adapting to climate change.

Liechtenstein , is highly dependent on the stability of its ecosystems as it is a mountainous country. This has strongly motivated the introduction of an active national climate policy and the participation of the country in international processes, such as the Alpine Convention. Strategies to address climate change as a new risk are largely of a regulatory nature and are all designed primarily to address the issues related to sustainability, i.e., by introducing sector-oriented legal documents, such as the Nature and Landscape Protection Act (1996); the Forest Act (1991); the Preservation and Protection of Agricultural Lands Act (1992); the Ordinance on the integrated rehabilitation of the Alpine and mountain regions (1968) and the revised Tourism Act (2000).

In France , forest managers have been working on measures to improve the resilience of forests since the storms of December 1999 and the drought conditions associated with the 2003 heat wave. It is considered most important to develop a larger biodiversity among forest stands with more diverse varieties and through more genetic diversification. It is also necessary to install or regenerate species which are better adapted to present and future local conditions, so that they are more resilient to biotic and abiotic environmental conditions. Tools are being elaborated in order to improve the choice of species. More dynamic forest management practices with wider spacing and strong and early thinning can reduce vulnerability to wind storms and at thesame timeimprove thewater budget andtherefore resistance to drought conditions. Better forest management practices and the presence of understory vegetation increases biodiversity and soil protection improves recycling of mineral elements and reduces mineral leaching. Development of heating plants or district heating using wood has been proposed in order to mitigate climate change. This would also help to utilise forest residues after storms and decrease the regeneration cost of damaged forests.

As a means to implement the Habitats Directive 92/43/ EEC, the German Federal Nature Conservation Act (of 25 March 2002) states that the Federal Laender shall establish a network of interlinked biotopes covering at least 10% of the total area of each Federal Land. The required core areas connecting areas and connecting elements shall be legally secured via the designation of appropriate areas, detailed planning in accordance with the provisions of planning law, long-term arrangements (contractual nature conservation) or other appropriate measures. This is intended to safeguard an interlinked network of biotopes in a sustainable fashion. Such a network of interlinked biotopes is particularly important and represents a dynamic response of ecosystems to global change to protect biodiversity.

In their national communications, many countries reported their concerns over issues such as goods and services of terrestrial ecosystems (e.g., timber production, biodiversity etc.) that may be threatened under a changing climate. But very few countries go beyond the list of general adaptation options for forest management and biodiversity protection. Emerging from this is the inadequacy of knowledge on potential impacts in terrestrial ecosystems and practical guidance for adaptation.

In addition, some multi lateral initiatives have been taken in order to establish a stronger (i.e., ‘climatically robust’) network of ecological areas within Europe. An example of such an initiative is the Pan-European Ecological Network PEEN.

4.4.2 Managing Europe's water resources

Rising temperature and changing rainfall patterns are expected to change the availability of water resources in Europe.

The integrated management plans for water resources in Spain constitute one component for adaptation to climate change. The National Water Plan, Law 10/2001 of 5 July (Analysis of Water Systems) accounts for potential climate change induced reduction in water availability and analyses the effect of these reductions on management and planning.

Drought mitigation has been recognised as a national priority for Hungary. The improvement of drought forecast, for example, through development of reliable drought indices is recommended as an important adaptation measure. A national drought mitigation strategy is to be developed.

In Greece , an integrated water management plan is considered imperative to address the present day problem of low water use efficiency, which results from the combination of irrigation and cultivation practices. The preparation for the full implementation of this plan has started. Cross governmental departments have drafted a legislative framework for its implementation.

In the Netherlands , climate change and adaptation measures are explicitly integrated into the water policy agenda. Emphasis is placed on ‘no-regret’ strategies. Although flood risks seem to dominate the adaptation agenda in water policy, the increased risk of dry spells and water shortage are also recognised. The spatial implications of the Cabinet's position on water management and the associated adaptation measures have been incorporated in the Dutch Spatial Policy. Inclusion in the policy for Rural Areas offers an opportunity to combine the implementation of measures in rural areas for increased safety and flood prevention with measures for such objectives as improving water quality, combating dropping water-tables, reconstructing rural areas and improving the ecological infrastructure.

Organizations involved in providing water services also started to explore the implications of climate change in terms of vulnerability and options for adaptation. The Norwegian Water Resources and Energy Directorate is an example. In the United Kingdom , organizations in the water resources sector are also taking actions to prepare for a changing climate.

4.4.3 Protecting people and infrastructure from coastal and river floods

Coastal and low-lying areas constitute a substantial part of Europe. With changing rainfall pattern (including extremes) and global warming induced sea level rise, a number of countries will be facing increased risk of coastal and river flooding. Benefiting from the long tradition of dealing with extreme weather events, flood defence is among the areas with best developed adaptive measures. Policies, guidance documents, regulations, and even concrete technical adaptation actions have been developed at the EU, national, and sub-national levels. Some of these measures are not deliberately designed for adaptation to long-term climate change impacts, though. Instead, they are developed for addressing short-term extremes.

At EU level, a flood prevention and management action plan is being developed. EU Environment Ministers asked the European Commission to table a formal proposal for the plan, which will be based on solidarity and will make provision for an early warning system, integrated flood basin and flooding management plans and the development of flood risk maps. The plan includes a possible future Floods Directive.

At national level, many northern and western European countries have national flood management policies and guidelines. Integrating new information on climate change and its potential impacts, such policies and guidelines are being reviewed and adjusted periodically.

In the United Kingdom , the trauma of human misery and property loss caused by the 1953 coastal flooding alerted the Government to the potential dangers. A radical rethinking led to major new flood defence infrastructure being built and eventually the commissioning of the Thames Barrier in 1987. The first major IPCC Assessment (1991) led to changed approaches to coastal planning throughout the United Kingdom. This has incorporated an allowance for climate change and sea level rise built into all new coastal flooding infrastructure. Planning Policy Guidance 25, published by the Office of the Deputy Prime Minister (ODPM, 2001), takes a precautionary approach to managing development and flood risk. It aims to direct new development away from areas at highest risk of flooding and takes account of climate change….

In the Netherlands , criteria and boundary conditions for the safety features of all dykes and other protection infrastructure are periodically updated to incorporate the available information on climate change and other environmental changes. The Dutch coastal policy plan (3rd Coastal Policy, 2000) strongly emphasises the new challenges caused by climate change, especially sea level rise and an increase in the number of storms. Various national and sub-national policy plans like the Dutch Spatial Policy (Ministry of Housing, 2004) and the Water Policy Plan 21st Century (Nota Waterbeleid 21ste eeuw) (Ministry of Transport, 2000) recognize the need for adaptation in water management and coastal zone management. An example of an adaptation action for rivers is in Hengelo, where the peak flow of the Woolderbinnenbeek can be reduced by 60%, to prevent the downstream agricultural land and town centres from getting flooded.

european environment agency (eea).”vulnerability and adaptation to climate change in europe” (2005)<> (accessed november 21, 2007).

See Also Coastal Populations; Coastlines, Changing; Economics of Climate Change; Energy Efficiency; Extinction; IPCC Climate Change 2007 Report: Impacts, Adaptation and Vulnerability; Lifestyle Changes; Refugees and Displacement; Social Cost of Carbon (SCC).



Parry, M. L., et al, eds. Climate Change 2007: Impacts, Adaptation and Vulnerability: Contribution of Working Group II to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. New York: Cambridge University Press, 2007.


Davis, Margaret B., et al. “Range Shifts and Adaptive Responses to Quaternary Climate Change.” Science 292 (2001): 673–679.

deMenocal, Peter B., et al. “Cultural Responses to Climate Change During the Late Holocene.” Science 292 (2001): 667–674.

Giles, Jim. “How to Survive a Warming World.” Nature 446 (2007): 716–717.

Kabat, Pavel, et al. “Climate Proofing the Netherlands.” Nature 438 (2005): 283–284.

Parmesan, Camille, and Gary Yohe. “A Globally Coherent Fingerprint of Climate Change Impacts Across Natural Systems.” Nature 421 (2003): 37–42.

Revkin, Andrew C. “Aid to Help Asia and Africa with Effects of Warming.” The New York Times (August 9, 2007).

Web Sites

“China Now No. 1 in CO2 Emissions; USA in Second Position.” Netherlands Environmental Assessment Agency, June 22, 2007. <> (accessed November 10, 2007).

“Strategies for Adaptation to Sea Level Rise.” Intergovernmental Panel on Climate Change, Response Strategies Working Group, Coastal Systems Subgroup, 1990.<$File/adaption.pdf> (accessed November 10, 2007).

Larry Gilman

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Adaptation is a term used to describe the ways in which organisms change over time in response to the changing demands of their environment. Organisms seem to accumulate certain physiological, behavioral, and structural traits gradually, and these traits aid them in their ability to survive and reproduce under existing environmental conditions. The grasping hands of primates, the sensitive antennae of insects, and the flowers and fruits of plants are all forms of adaptation that promote survival, reproduction, or both.

Historical background

Up until the eighteenth century, scientists generally believed that every species was created separately and remained unchanged for centuries. Many features of living thingsthe bee's sting, the vertebrate (an animal with a backbone) eye, the human brainappeared to have been designed by a master engineer to serve their specific purpose. A philosophy known as natural theology, which arose in the seventeenth century, argued that the elegant and often complex features of organisms were the products of a direct design by God.

But during the eighteenth century, the scientific community began to take a closer look at the immense diversity (the vast differences) and interrelatedness of (connections between) living things. The excavation of plant and animal fossils prompted a new view that life on Earth developed gradually and unevenly from simple to advanced organisms. The observation that species have "adapted" to survive in particular habitats raised new questions about the ways organisms could "fine-tune" themselves to meet the demands of their environment.

Words to Know

Adaptation: From the Latin ad ("toward") plus aptus ("fit for some role"); any structural, physiological, or behavioral trait that aids an organism's survival and ability to reproduce in its existing environment.

Coadaptation: Mutual dependence between members of two species.

Evolution: The theory that all plants and animals developed gradually from earlier forms over a long period of time and that variations within a species are the result of adaptive traits passed on from generation to generation.

Exaptation: Any adapted trait that performs a beneficial function different from the one it originally evolved to serve.

Natural selection: A natural process that results in the survival of individuals or groups best adapted to the conditions in which they must exist.

Different species that live in different environments often exhibit similar characteristics. Fish, whales, and penguins all use fins or flippers to propel themselves through water. The common plan of such features demanded an explanation, so researchers set out to explain this similarity of traits in unrelated organisms.

The Peppered Moth

Henry Bernard David Kettlewell's study of the peppered moth, Biston betularia, is one of the most widely cited cases of natural selection producing adaptation. Before the Industrial Revolution in England in the late 1800s, this moth was predominantly light in color. (The Industrial Revolution was a time of major change in England's economy, marked mainly by the introduction of power-driven machinery.) Light coloring afforded perfect camouflage for the moth from predatory birds, since it blended so well with the similarly colored lichen-covered tree trunks on which it rested. When pollution from factories caused the lichen on the trees to die, the moths' resting place became the darker color of the bark beneath. Kettlewell observed that, as this environmental transformation occurred, a dark form of the moth became increasingly common, eventually making up more than 90 percent of the population of moths in the affected areas. In the unpolluted areas, however, the original light form of the moth remained common.

Kettlewell attributed the moth's color change to selection by predatory birds, which locate the moths by sight, and so remove (by eating) individual moths that do not blend in with the background coloration of trees in their environment. He tested his idea that the moths' color protected them from predation (being captured) by placing each of the two forms on trees in different areas, photographing birds in the act of capturing moths, and measuring the rates at which the two moth forms were eaten by birds. Kettlewell concluded that the moths' color was indeed the result of adaptation to conditions in their habitat.

One effort to explain adapted traits was proposed by French botanist Jean-Baptiste Lamarck (17441829). In 1809, he theorized that changes in the environment cause structural changes during an organism's life that are passed on to offspring. According to the notion of the inheritance of acquired characteristics, also known as Lamarckism, giraffes would have "acquired" their long necks from stretching to reach leaves not available to other animals. Members of each succeeding generation stretched their necks to attain leaves at a higher level, which led to the modern giraffe. Although Lamarck's theory was later discredited, he remains the first scientist to acknowledge the adaptability of organisms.

In 1859, Charles Darwin (18091882), the great English naturalist, published his influential book The Origin of Species by Means of Natural Selection. In it, Darwin discusses the adaptations of organisms as the product of natural selection. Natural selection implies thatwhen forced to compete for limited resources such as foodthose organisms best adapted to their specific environment are most likely to survive, reproduce, and transmit their traits to offspring.

Biologists now recognize natural selection as the means by which evolutionary adaptation occurs. Exactly how specific adaptations arise, however, is far from solved. It is easy to imagine how natural selection might produce relatively simple adaptations such as camouflage: a rabbit that lives in regions covered by snow in winter is better protected from prey if it produces a white coat during the winter months. The difficulty arises in explaining the evolution of extremely complex adaptations, such as the vertebrate eye. Invertebrates, animals lacking a backbone, have simple eyes that detect only changes in light and form only a poor image at

best. Vertebrates, animals with a backbone, detect changes in light and motion and can form detailed images. How did such a superb adaptation come about?

Darwin suggested the answer lies in very gradual changes over many generations, in which each intermediate stage leading to a fully formed eye had some adaptive value. All the parts making up a fully functioning eye could evolve independently in small steps, each one building on and interacting with earlier changes. Thus, even a partially developed eye could be quite advantageousindeed, could mean the difference between life and deathfor an ancient vertebrate.

Examples of adaptation

Physiological adaptation. People who visit or live at high altitudes undergo physiological changes (adaptations) to adjust to the low-oxygen environment. Travelers to these areas commonly experience hypoxia, a condition of low oxygen in the blood. To compensate for the temporary drop in oxygen, vacationers' bodies speed up the oxygenation process: they breathe at a faster rate, their hearts speed up and pump more strongly to send more blood throughout the body, and they produce more red blood cells to carry oxygen to body tissues. Over a longer period, as the body adjusts to the change in altitude, the heart output and ventilation rate return to normal levels, but the red blood cell count continues to climb. The most famous of all high-altitude peoples are the Sherpas of Nepal, whose climbing feats offer a stunning example of evolved adaptation.

Evolutionary adaptation. Some of the most interesting cases of adaptation occur when two species evolve together so that each benefits from the other. This mutual dependence can be seen between the ant Formica fusca and the larval (not yet fully developed) stage of the lycaenid butterfly Glaucopsyche lygdamus. The butterfly caterpillar produces a sweet "honeydew" solution that the ants harvest as food. In return, the ants defend the caterpillar against parasitic wasps and flies. The mutual adaptation of two species in this manner is known as coadaptation.

Interactions between species are not always beneficial for both members, however. Heliconius butterflies scatter the pollen from the flowers of Passiflora vines, benefitting the plant. But female butterflies also lay single eggs on young Passiflora shoots, and the developing larva may eat the entire shoot, a definite cost to the plant. As an apparent adaptive response, several Passiflora species produce new shoots featuring a small structure that closely resembles a Heliconius egg. A female butterfly that sees this "egg" will avoid laying her own egg there, and the shoot will be spared.

Current approaches to adaptation

Some researchers draw a distinction between current use and the historical origin of adaptive traits. This distinction has led researchers Stephen Jay Gould and Elisabeth Vrba to suggest the term exaptation to describe traits that were originally intended to perform different functions in an organism. For instance, the observation that a white coat helps camouflage a snowshoe hare in the snow does not provide information about the origin of the species' white fur. It may have evolved for its improved heating properties and only by chance proved to be advantageous as camouflage. In some cases, therefore, the features we recognize as adaptive are really only secondary uses of traits that originally arose for other reasons.

[See also Evolution ]

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To survive and reproduce, all living organisms must adjust to conditions imposed on them by their environments. An organism's environment includes everything impinging upon it, as well as everything that is affected by that organism. Conformity between an organism and its environment constitutes what biologists call adaptation.

Biotic and Abiotic Environments

Plants and animals have adapted to their environments genetically and by means of physiological, behavioral, or developmental flexibility, including both instinctive behavior and learning. Adaptation has many dimensions in that most organisms must conform simultaneously to numerous different aspects of their environments. Adaptation involves coping not only with the physical abiotic environment (light, dark, temperature, water, wind), but also with the complex biotic environment (other organisms such as mates, competitors, parasites , predators, and escape tactics of prey). Conflicting demands of these various environmental components often require that an organism compromise in its adaptations to each.

Conformity to any given dimension requires a certain amount of energy that is then no longer available for other adaptations. The presence of predators, for example, may require that an animal be wary, which in turn is likely to reduce its feeding efficiency and hence its competitive ability.

For a small bird, trees are an important part of its environment: They offer vital shade during the heat of a hot summer day, places to forage for insects, safety from ground-dwelling predators, and safe places to build nests and raise chicks. Blades of grass or hairs used to line a bird's nest are also important components of a bird's environment. During the dangerous night, a bird copes with nocturnal predators such as raccoons by sleeping perched on a small twig high above the ground. While gleaning tiny insects from tree leaves during the day, a bird remains alert for diurnal predators like hawks.

Many birds cope with changing seasonal conditions by migrating to warmer places at lower latitudes where there is more food. Over eons of time, natural selection has molded birds to make them effective at escaping from the predictable dire consequences of winter (a time of high mortality). Birds that did not successfully evade winter's icy clutches died without leaving any surviving offspring, whereas those that migrated survived to pass on their genes. Natural selection has endowed birds with a built-in biological clock, which they compare against day length, effectively giving them a builtin calendar. Changing day length affects a bird's pituitary gland, causing it to secrete hormones that control avian behavior. Short autumn days elicit a "wanderlust," ultimately leading to migratory behavior. Experiments with migrating birds in planetaria have shown that tiny bird brains have been hard-wired so that they contain a map of the stars. Indeed, natural selection "invented" celestial navigation.

Factors that Affect Adaptation

Organisms can conform to and cope with a highly predictable environment relatively easily, even when it changes in a regular way, as long as the changes are not too extreme. Adaptation to an unpredictable environment is usually more difficult; adapting to extremely erratic environments may even prove impossible. Many organisms have evolved dormant stages that allow them to survive unfavorable periods, both predictable and unpredictable. Brine shrimp in deserts and annual plants everywhere are good examples. Brine shrimp eggs survive for years in the salty crust of dry desert lakes; when a rare desert rain fills one of these lakes, the eggs hatch, the shrimp grow rapidly to adults, and they produce many eggs. Some plant seeds known to be many centuries old are still viable and have been germinated.

Very small undirected changes in the physical environment can sometimes improve the level of adaptation between an organism and its environment, but large changes are almost always detrimental. Changes in the environment that reduce overall adaptation are collectively termed the "deterioration of environment." Such changes cause directional selection resulting in accommodation to the new environment, or adaptation. Changes in biotic environments (such as the hunting efficiency of an organism's predator) are usually directed and typically reduce the level of adaptation.

Every individual is simultaneously a member of a population, a species, and a community; therefore, it must be adapted to cope with each and must be considered in that context. An individual's fitnessits ability to perpetuate itself as measured by its reproductive successis greatly influenced by its status within its own population. An individual might be a resident or a vagrant, mated or unmated, or high or low in a pecking order, all factors that strongly affect its fitness. Any given individual's fitness is also influenced by various interspecific associations of its species and especially by the particular community in which it finds itself embedded.

"Arms Races"

Individuals and species must "track" their environments in ecological and evolutionary time, adapting and evolving as their environments change. Natural selection acting on natural enemies (prey, parasites, and predators) will always result in a deterioration of an organism's biotic environment, diminishing fitness. Every prey-predator or host-parasite interaction constitutes an escalating "arms race," in which moves alternate with countermoves.

Prey that are better able to escape from their predators, or hosts that can better resist infection by parasites, will enjoy a fitness advantage. But better predators and better parasites are also favored by natural selection themselves, assuring that the arms race will continue to escalate indefinitely. Indeed, most species are probably evolving rapidly just to maintain a given current level of adaptation in the face of a continually deteriorating environment. Still other interactions between species are mutually beneficial, resulting in increased fitness for both parties, such as between plants and their pollinators.

Any genetically based physiological, behavioral, or ecological trait that enables an organism to cope with, and to survive and reproduce in, its environment represents an adaptation. Some traits may not be adaptive but simply leftover vestiges of traits that once were adaptive. A given trait can also be "preadapted" if it was formerly adaptive under some prior set of conditions now gone but is later co-opted as the basis of a new adaptation under some new environmental conditions. For instance, it is likely that bird feathers were initially important for temperature regulation, rather than for flying.

see also Community; Convergent Evolution; Evolution; Natural Selection; Parasitic Diseases; Pituitary Gland; Population Dynamics; Predation and Defense; Sexual Selection; Symbiosis

Eric R. Pianka


Fisher, Ronald A. The Genetical Theory of Natural Selection. Oxford: Clarendon Press, 1930.

Pianka, Eric. R. Evolutionary Ecology, 6th ed. San Francisco: Addison-Wesley-Longman, 2000.

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The act or process of modifying an object to render it suitable for a particular or new purpose or situation.

In the law of patents—grants by the government to inventors for the exclusive right to manufacture, use, or market inventions for a term of years—adaptation denotes a category of patentable inventions, which entails the application of an existing product or process to a new use, accompanied by the exercise of inventive faculties. Federal law provides: "Whoever invents or discovers any new and useful process, machine, manufacture, or composition of matter, or any new and useful improvement thereof, may obtain a patent therefore, subject to the conditions and requirements of this title." 35 U.S.C.A. §101.

The adaptation of a device to a different field can constitute an invention if inventiveness exists in the conception of new use and with modifications necessary to render the device applicable in the new field. The progressive adaptation of well-known devices to new, but similar, uses is merely a display of an expected technical proficiency, which involves only the exercise of common reasoning abilities upon materials furnished by special knowledge ensuing from continual practice. It, therefore, does not represent a patentable invention. Ingenuity beyond the mere adaptation of teachings as could be done by a skilled mechanic is required to achieve a patentable invention; inventive talent, rather than skill in adaptation, must be manifested. To entitle a party to the benefit of the patent statute, the device must not only be new; it must be inventively new. The readaptation of old forms to new roles does not constitute invention where there is no significant alteration in the method of applying it or in the nature of the result obtained. No invention will be recognized if the new form of the result has not previously been contemplated and, irrespective of the remoteness of the new use from the old, if no modifications in the old device are necessary to adapt it to the new use.

Invention is generally not involved where an old process, device, or method is applied to a new subject or use that is analogous to the old or to a new use or the production of a new result in the same or analogous field. If the new use is so comparable to the old that the concept of adapting the device to the new use would occur to a person proficient in the art and interested in devising a method of changing the intended function, there is no invention even though significant alterations have been made. The application of an old device to a new use is normally patentable only if the new use is in a different field or involves a completely novel function. In addition, the physical modifications need not be extensive, as long as they are essential to the objective.

In the law of copyrights the exclusive right of the author of a literary project to reproduce, publish, and sell his or her work, which is granted by statute, adaptation refers to the creation of a derivative work, which is protected by federal copyright laws.

A derivative work involves a recasting or translation process that incorporates preexisting material capable of protection by copyright. An adaptation is copyrighted if it meets the requirement of originality, in the sense that the author has created it by his or her own proficiency, labor, and judgment without directly copying or subtly imitating the preexisting material. Mere minor alterations will not suffice. In addition the adapter must procure the consent of the copyright owner of the underlying work if he or she wants to copy from such work. The copyright in a derivative work, however, extends only to the material contributed by the adapter and does not affect the copyright protection afforded to the preexisting material.

The rise in the use of digital media has caused new dilemmas in the area of copyright law with respect to adaptations. Even average technology users may make copies and adapt the original works to their needs. Recent issues in this area have focused upon intellectual property rights in the context of the internet and computer programs.

Even average computer users are now capable of copying digital music files and modifying them through the use of software. The Internet now allows these users to prepare these modifications and distribute them to a wide audience using the Web, e-mail, and other methods of distribution. The Copyright Act of 1976 continues to protect the copyright holders, generally requiring those who prepared derivative works to obtain permission from the copyright holder (17 U.S.C.A. § 114(b) [1996]). However, enforcement of these provisions has proven difficult and led to a number of efforts, including those by the Recording Industry Association of America, to find new methods for protecting the rights of the copyright holders.

A second cause of concern among copyright owners is the ability of computer users to make copies of computer program and adopt these programs to serve the users' purposes. The Copyright Act provides an exclusive right to the copyright holders of computer programs and allows owners of copies of these programs to make additional copies only in limited circumstances (17 U.S.C.A. § 117 [1996]). Like sound recordings, protection of these copyrights has proven difficult, leading lawmakers to consider a number of new options to protect these rights.

In the law of real property, with respect to fixtures (articles that were personal property but became part of the realty through annexation to the premises), adaptation is the relationship between the article and the use that is made of the realty to which the article is annexed.

The prevailing view is that the adaptation or appropriation of an article affixed to real property for the purpose or use to which the premises are devoted is an important consideration in ascertaining its status as a fixture. According to this theory, if the article facilitates the realization of the purpose of the real property, the annexor presumably intends it to be a permanent accession. Numerous other cases, however, allude to the adaptation of an item to the use to which the premises are designated, as merely one of the tests or factors that should or must be evaluated in determining that it constitutes real property. Other cases view the character of the use of the article annexed as significant.

The special construction or fitting of an article for location and use on certain land or in a particular building, which mitigates against use in another location, indicates that is was intended to constitute a part of the land.

The adaptability of an annexed article for use in another location is sometimes viewed as demonstrating the retention of its character as personalty (personal property), but this characteristic is not conclusive. Articles not designed to comprise the realty retain their character as personalty.

further readings

Benn, Marvin N., and Richard J. Superfine. 1994. "§ 117—The Right to Adapt into the Fourth Generation and the Source Code Generator's Dilemma." John Marshall Journal of Computer and Information Law 537.

Miller, Arthur R., and Michael H. Davis. 2000. Intellectual Property: Patents, Trademarks, and Copyright in a Nutshell. 3d ed. St. Paul, Minn.: West Group.

Plotkin, Mark E., ed. 2003. E-Commerce Law & Business. New York: Aspen.

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Adaptation is not part of Freudian vocabulary (it does not appear in the index of the Standard Edition, for example). The idea of adaptation, however, is present throughout Freud's work. It appears as early as 1895, in his "Project for a Scientific Psychology" (1950a), when he discusses the mechanisms of perception, attention and memory. The idea runs through all of Freud's subsequent work whenever he discusses the relation between psychic reality and the "reality of the outside world." It is found, for example, in "Instincts and their Vicissitudes" (1915c) and "Repression" (1915d), when he writes that dangers that can't be avoided through behavioral means are "rejected toward the interior." Other texts where the concept appears include "Neurosis and Psychosis" (1924b), "The Loss of Reality in Neurosis and Psychosis" (1924e), and "An Outline of Psycho-Analysis" (1940a). In fact, there are few texts by Freud where the question of adaptation isn't found, even if the word itself rarely appears.

Adaptation and the related theoretical issues are central to the development of ego-psychology, which was, for the most part, based on Freud's structural theory and the work of Anna Freud (1936/1937) and Heinz Hartmann, author of Ego Psychology and the Problem of Adaptation (1938/1958). It was in this period that a theorical schism developed, leading to differences in clinical psychoanalytic practice between those analysts (especially English-speaking) who adapted this point of view and those who preferred other options, either along the lines developed by Melanie Klein and her successors or the rather different approach taken by Lacan and his successors.

Jacques Lacan was, in fact, highly critical of the primacy given to the problems of adaptation in ego-psychology. He emphasized that naively establishing "external reality" as a given prior to and outside of psychic activity is a theoretical absurdity since that exterior reality is constructed through close interaction with psychic reality itself. He also pointed out the dangers of an analytical practice in which the analyst, within the framework of a normative and "normalizing" enterprise, developed mastery, or even a sense of excessive power, in assuming that his or her own "adaptation" is by definition better than that of the patient. Whatever one might think of these criticisms and their rebuttals, there is little doubt that they have had considerable impact, well beyond the field of Lacanian thought, especially in the French-speaking world. Unfortunately, this has had the effect of "throwing the baby out with the bathwater" through the unjustified condemnation of any psychoanalytic consideration of the problems of adaptation. These problems cannot be avoided, however, to the extent that psychic processes are constantly being adjusted in terms of their internal equilibrium and modified as a result of the impact of outside events.

Roger Perron

See also: Defense; Ego; Ego Psychology and the Problem of Adaptation ; Individuation (analytical psychology); Kardiner, Abram; Normality; Pichon-Rivière, Enrique; Self (true/false).


Canguilhem, Georges. (1989). The normal and the pathological (Carolyn R. Fawcett & Robert S. Cohen, Trans.). New York: Zone Books. (Original work published 1966)

Freud, Anna. (1937). The ego and the mechanisms of defense. London: Hogarth Press. (Original work published 1936)

Freud, Sigmund. (1915c). Instincts and their vicissitudes. SE, 14: 109-140.

. (1915d). Repression. SE, 14: 141-158.

. (1924b [1923]). Neurosis and psychosis. SE, 19: 147-153.

. (1924e). The loss of reality in neurosis and psychosis. SE, 19: 180-187.

. (1940a [1938]). An outline of psycho-analysis. SE, 23: 139-207.

. (1950a [1887-1902]). Extracts from the Fliess papers. SE, 1: 173-280.

Hartmann Heinz. (1958). Ego psychology and the problem of adaptation (David Rapaport, Trans.). New York: International Universities Press. (Original work published 1938)

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Adaptation refers to any change that makes a species or an individual better suited to its environment or way of life. All living creatures must be able to adapt to changes in their environment if they are to survive and reproduce. The process of adaptation does not always result in an obvious physical change, but may affect an individual's behavior or even its internal processes.

Adaptation could be described as the theory of evolution in action. This theory was first offered in its best and most complete form in 1859 by the English naturalist, Charles Darwin (1809–1882). Darwin's theory of evolution suggested that all living things are subject to a gradual process of change over a long period of time. Evolution is therefore the process that results in living things changing through successive generations. Darwin also described a mechanism called "natural selection," which is the means through which these hereditary changes are passed on from one generation to another.

Darwin's theory of evolution by natural selection—which is highly important to understanding the life sciences—states that individual organisms possessing certain traits or characteristics that are most suited to a particular environment have a better chance of surviving and therefore of passing these traits on to their offspring. In other words, organisms in possession of favorable traits allows these traits to be "selected" by nature (natural selection) so that these organisms survive and produce young that have the same favorable traits.

Darwin's years of travel, study, and thought led him to make three important observations or conclusions. First, he stated that all living things vary, or that each individual varies slightly from the others of its species. This could be seen in any given species in which differences could be found among any group of the same organisms (such as a flower with an extra petal or a deer with larger-than-average antlers). Second, Darwin suggested that individuals were able to pass certain characteristics on to their young, who inherited them. Modern genetics has proven this to be true. Third, Darwin noted that all life is involved in a struggle for survival. To Darwin, these three observations explained why nature allowed most organisms to produce far more offspring than could ever survive. Offspring that did survive and were able to reproduce, according to Darwin, were usually the ones who possessed certain traits better suited to their environment. These individuals had a better chance of surviving and of producing more individuals like themselves. Over many generations, it was simply nature's "selection" of the individuals with the fittest (or best adapted) characteristics that explained Darwin's theory of evolution. This accounts for the well-known phrase, "survival of the fittest."

"Survival of the fittest" could also be described as "survival of the best adapted." Among any group of individuals or organisms of the same species, there will always be variations or differences (in color, shape, behavior, and even chemical makeup). An adaptation then, is considered to be any variation that makes an organism better suited to its environment. Camouflage is one way nature has of providing protection for an individual, and an organism whose color or shape allows it to blend into its environment is more likely to survive and reproduce than one whose coloration makes it easier to be noticed. Woodpeckers are highly specialized birds and are a good example of the process of adaptation. Their main job is to find and eat insects that live in and beneath the bark of trees. Consequently, those woodpeckers with the most powerful and chisel-like beaks, strong neck muscles for hammering, sturdy skulls, grasping feet, stiff-supporting tail feathers, and long tongues proved best able to survive and pass on these traits.

Darwin first described his theory of evolution by natural selection in 1859 in his classic book, On the Origin of Species by Means of Natural Selection. The idea of adaptation is key to Darwin's theory. Darwin distinguishes between genotypic adaptation and phenotypic adaptation. The type of adaptation discussed above—in which an individual possesses a "favorable" gene for a certain characteristic which it passes on to its offspring—is called genotypic or evolutionary adaptation. This type of adaptation is genetic, permanent, and is very different from phenotypic adaptations. In contrast to genotypic adaptations, phenotypic adaptations are traits that are developed during an individual's lifetime. An example of phenotypic, or nongenetic, adaptation might be a certain type of behavior that is learned or developed by an individual. The macaque monkeys in Japan, for example, have learned to wash their food in water, and newborns soon copy this behavior. Although it is not known exactly why the first macaque washed its food, this behavior in not instinctive with them. Rather it is a case of learned behavior.

[See alsoEvolution; Evolution, Evidence of; Evolutionary Theory ]

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An adaptation is any developmental, behavioral, physiological, or anatomical change in an organism that gives that organism a better chance to survive and reproduce. The word adaptation also refers to the fitting of a whole species, over time, to function in its particular environment, and to those specific features of a species that have changed.

At the level of the individual organism, an adaptation is a change in response to conditions. This is a short-term change with a short-term benefit. Adaptations acquired by individuals during their lifetime, such as muscles strengthened by exercise or behaviors honed by experience, make an individual organism better adapted. Species as a whole, however, generally become better adapted to their environments only by the process of natural selection. Except in the form of learned behavior, adaptations achieved by individual organisms cannot be passed on to offspring.

Less-adapted species are less perfectly attuned to a particular environment but may be better-suited to survive changes in that environment or to colonize new areas. Highly adapted species are well suited to their particular environment, but being more specialized, are less likely to survive changes to that

environment or to spread to other environments. An example of a highly adapted species would be a flower that depends on a specific insect that exists only or primarily in its present environment for pollination. The plant may achieve highly reliable pollination by these means, but if its target species of insect becomes extinct, the plant will also become extinct unless the species can adapt to make use of another pollinator.

Microorganisms are capable of adaptation. A classic example is the response of some bacteria to increased temperature. This response includes the production of a variety of proteins known as heat shock proteins that function to increase the stability of the membrane(s) that hold the cell together, aid in the transport of compounds into and out of the cell, and other functions that increase the likelihood of the cells survival under the more adverse conditions.

Another type of adaptation is sensory adaptation. If a receptor or sense organ is over-stimulated, its excitability is reduced. For example, continually applied pressure to an area of skin eventually causes the area to become numb to feeling and a considerably larger pressure has to be applied to the area subsequently to elicit a similar response. This form of adaptation enables animals to ignore most of their skin most of the time, freeing their attention for more pressing concerns.

Whether occurring within a span of minutes, over an organisms lifetime, or over thousands or millions of years, adaptation serves to increase the efficiency of organisms and thus, ultimately, their chances of survival.



Gould, Stephen Jay. The Structure of Evolutionary Theory. Cambridge, MA: Harvard University Press, 2002.

Hochachka, Peter W., and George N. Somero. Biochemical Adaptation: Mechanism and Process in Physiological Evolution. New York: Oxford University Press, 2002.

Storey, Kenneth B. Functional Metabolism: Regulation and Adaptation. New York: Wiley-Liss, 2004.

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adaptation, in biology, has several meanings. It can mean the adjustment of living matter to environmental conditions and to other living things either in an organism's lifetime (physiological adaptation) or in a population over many many generations (evolutionary adaptation). The word can also refer to a trait that is considered an adaptation. The ability to adapt is a fundamental property of life and constitutes a basic difference between living and nonliving matter.

Most living things require free oxygen from the air or from water, but yeasts, many bacteria, and some other simple forms obtain the oxygen required for oxidation from molecules of substances that contain the element. Various animals and plants are adapted for securing their food and for surviving the extremes of temperature and of water supply in desert, tropical, and polar regions. For most organisms the optimum temperature is between about 20°C (68°F) and 40°C (104°F). Some algae and protozoans live in hot springs, and some bacteria can survive freezing or survive on chemicals, without light, in the ocean depths. Cacti can survive heat and drought. Certain fish and other aquatic animals live in deep water and are so specialized to withstand the great pressure that they burst if lifted to sea level.

Animals show anatomical adaptations—e.g., the body of the fish is suited to life in the water; the body of the bird is adapted for flight; and the land mammals show a wide variation in the structure of limbs and body that enables some to run swiftly, some to climb, some to swing from tree to tree, some to glide through the air, and others to jump. The whale, an aquatic mammal, can adjust to great pressure changes at different levels in the water. The beaks of birds vary in shape and size according to what they feed on—e.g., on seeds, on insects, on aquatic animals, or on small mammals. The feet and legs of birds also show modifications that fit them for perching, for wading, or for paddling through the water. Adaptive coloration is observed in many animals (see protective coloration). Among communal insects, such as ants and honeybees, the individuals are highly adapted to perform their functions in the community.

It is believed by many scientists that life originated in the sea and that through gradual evolutionary changes some forms became adapted to life on land. Variations may arise as a result of mutation, or of recombinations of the genes in the germ cells. Such variations are inherited (see genetics). Those that aid the organism to meet the conditions of a changing environment or help it in its competition with other living things enable it to survive and reproduce, the changes thus being passed on from one generation to another and in this way perhaps producing a new species.

See ecology; evolution; selection.

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Adaptations are the most important features of all organisms. An adaptation is a trait or characteristic that makes an animal survive or reproduce better in its environment. These traits can be morphological , physiological , or behavioral . Adaptations include almost all kinds of traits, such as what makes an organism blend into its surroundings, find food, mate with the correct species, and be able to survive.

An example of a morphological adaptation that increases the chance of survival is the coloration of an animal. Most animals that live in the arctic snow are white. Being white helps them blend in with the snow and hide from predators. An example of a physiological adaptation that increases the likelihood of survival is the kangaroo rat's metabolism. Kangaroo rats live in the desert of the North American Southwest. It is extremely hot and dry there, and very little water is available. The kangaroo rat never needs to drink water because its metabolism has changed and adapted to conserve water; it gets all the water it needs from seeds it eats. Humans are the opposite; we must drink water daily because our metabolism uses lots of water. An example of a behavioral adaptation that increases reproduction is the croaking and calling of male frogs, which gets female frogs to come to the males for mating. Those frogs that call end up mating with more females and have more offspring than frogs that do not call.

Natural selection is the mechanism that produces adaptations. It is the difference in survival or reproduction between individuals with different traits. If an arctic hare, which lives in the snow, were gray rather than solid white, it would not survive as well as pure white hares. And if it does not live very long before it is killed by a predator such as a fox, then it will not produce as many offspring as a pure white hare, either. Natural selection increases the number of individuals in a population that actually has the adaptation; natural selection also maintains adaptations once animals have them.

All traits that are adaptations must have a genetic basis to them. That is to say, the trait must be produced in some way by genes , coded for by DNA. Genes enable an animal to pass on to its offspring the survival or reproductive advantage given by the adaptation.

There is one exception to the rule that adaptations need a genetic basistraits that are learned within an organism's lifetime. Something that is learned does not have a genetic basis; it was not inherited from the parents. An example of a learned adaptation is that some male songbirds learn the songs of their neighbors. Neighborhoods of birds sing similar songs, and females prefer males that know more songs. A male that has learned more songs will reproduce more; learning songs is an adaptation. But learned traits are not completely free from genetics; there is a genetic basis to the ability to learn, even if the actual learning does not have a genetic basis.

see also Biological Evolution; Morphological Evolution in Whales; Morphology; Natural Selection.

Laura A. Higgins


Campbell, Neil A., Jane B. Reece, and Lawrence G. Mitchell. Biology, 5th ed. Menlo Park, CA: Addison Wesley Longman, Inc., 1999.

Rose, Michael R., and George V. Lauder, eds. Adaptation. San Francisco, CA: Academic Press, 1996.