Adaptation

In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the population during that process. Thirdly, it is a phenotypic or adaptive trait, with a functional role in each individual organism, that is maintained and has evolved through natural selection.

Organisms face a succession of environmental challenges as they grow, and show adaptive plasticity as traits develop in response to the imposed conditions. This gives them resilience to varying environments.

History

Adaptation is an observable fact of life accepted by philosophers and natural historians from ancient times, independently of their views on evolution, but their explanations differed. Empedocles did not believe that adaptation required a final cause (~ purpose), but thought that it "came about naturally, since such things survived." Aristotle did believe in final causes, but assumed that species were fixed.[1]

Lamarck's Two-Factor Theory
The second of Jean-Baptiste Lamarck's two factors (the first being a complexifying force) was an adaptive force that causes animals with a given body plan to adapt to circumstances by inheritance of acquired characteristics, creating a diversity of species and genera.

In natural theology, adaptation was interpreted as the work of a deity and as evidence for the existence of God.[2] William Paley believed that organisms were perfectly adapted to the lives they led, an argument that shadowed Gottfried Wilhelm Leibniz, who had argued that God had brought about "the best of all possible worlds." Voltaire's Dr. Pangloss[3] is a parody of this optimistic idea, and David Hume also argued against design.[4] The Bridgewater Treatises are a product of natural theology, though some of the authors managed to present their work in a fairly neutral manner. The series was lampooned by Robert Knox, who held quasi-evolutionary views, as the Bilgewater Treatises. Charles Darwin broke with the tradition by emphasising the flaws and limitations which occurred in the animal and plant worlds.[5]

Jean-Baptiste Lamarck proposed a tendency for organisms to become more complex, moving up a ladder of progress, plus "the influence of circumstances," usually expressed as use and disuse.[6] This second, subsidiary element of his theory is what is now called Lamarckism, a proto-evolutionary hypothesis of the inheritance of acquired characteristics, intended to explain adaptations by natural means.[7]

Other natural historians, such as Buffon, accepted adaptation, and some also accepted evolution, without voicing their opinions as to the mechanism. This illustrates the real merit of Darwin and Alfred Russel Wallace, and secondary figures such as Henry Walter Bates, for putting forward a mechanism whose significance had only been glimpsed previously. A century later, experimental field studies and breeding experiments by people such as E. B. Ford and Theodosius Dobzhansky produced evidence that natural selection was not only the 'engine' behind adaptation, but was a much stronger force than had previously been thought.[8][9][10]

General principles

The significance of an adaptation can only be understood in relation to the total biology of the species.

— Julian Huxley, Evolution: The Modern Synthesis[11]

What adaptation is

Adaptation is primarily a process rather than a physical form or part of a body.[12] An internal parasite (such as a liver fluke) can illustrate the distinction: such a parasite may have a very simple bodily structure, but nevertheless the organism is highly adapted to its specific environment. From this we see that adaptation is not just a matter of visible traits: in such parasites critical adaptations take place in the life cycle, which is often quite complex.[13] However, as a practical term, "adaptation" often refers to a product: those features of a species which result from the process. Many aspects of an animal or plant can be correctly called adaptations, though there are always some features whose function remains in doubt. By using the term adaptation for the evolutionary process, and adaptive trait for the bodily part or function (the product), one may distinguish the two different senses of the word.[14][15][16][17]

Adaptation is one of the two main processes that explain the observed diversity of species, such as the different species of Darwin's finches. The other process is speciation, in which new species arise, typically through reproductive isolation.[18][19] A favourite example used today to study the interplay of adaptation and speciation is the evolution of cichlid fish in African lakes, where the question of reproductive isolation is complex.[20][21]

Adaptation is not always a simple matter where the ideal phenotype evolves for a given external environment. An organism must be viable at all stages of its development and at all stages of its evolution. This places constraints on the evolution of development, behaviour, and structure of organisms. The main constraint, over which there has been much debate, is the requirement that each genetic and phenotypic change during evolution should be relatively small, because developmental systems are so complex and interlinked. However, it is not clear what "relatively small" should mean, for example polyploidy in plants is a reasonably common large genetic change.[22] The origin of eukaryotic endosymbiosis is a more dramatic example.[23]

All adaptations help organisms survive in their ecological niches.[24] The adaptive traits may be structural, behavioural or physiological. Structural adaptations are physical features of an organism, such as shape, body covering, armament, and internal organization. Behavioural adaptations are inherited systems of behaviour, whether inherited in detail as instincts, or as a neuropsychological capacity for learning. Examples include searching for food, mating, and vocalizations. Physiological adaptations permit the organism to perform special functions such as making venom, secreting slime, and phototropism), but also involve more general functions such as growth and development, temperature regulation, ionic balance and other aspects of homeostasis. Adaptation affects all aspects of the life of an organism.

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

1. Adaptation is the evolutionary process whereby an organism becomes better able to live in its habitat or habitats.[25][26][27]
2. Adaptedness is the state of being adapted: the degree to which an organism is able to live and reproduce in a given set of habitats.[28]
3. An adaptive trait is an aspect of the developmental pattern of the organism which enables or enhances the probability of that organism surviving and reproducing.[29]

What adaptation is not

Tampering W Nature Guacharacas
Some generalists, such as birds, have the flexibility to adapt to urban areas.

Adaptation differs from flexibility, acclimatization, and learning. Flexibility deals with the relative capacity of an organism to maintain itself in different habitats: its degree of specialization. Acclimatization describes automatic physiological adjustments during life;[30] learning means improvement in behavioral performance during life.[31] These terms are preferred to adaptation for changes during life which are not inherited by the next generation.

Flexibility stems from phenotypic plasticity, the ability of an organism with a given genotype to change its phenotype in response to changes in its habitat, or to move to a different habitat.[32][33] The degree of flexibility is inherited, and varies between individuals. A highly specialized animal or plant lives only in a well-defined habitat, eats a specific type of food, and cannot survive if its needs are not met. Many herbivores are like this; extreme examples are koalas which depend on Eucalyptus, and giant pandas which require bamboo. A generalist, on the other hand, eats a range of food, and can survive in many different conditions. Examples are humans, rats, crabs and many carnivores. The tendency to behave in a specialized or exploratory manner is inherited—it is an adaptation. Rather different is developmental flexibility: "An animal or plant is developmentally flexible if when it is raised in or transferred to new conditions, it changes in structure so that it is better fitted to survive in the new environment," writes evolutionary biologist John Maynard Smith.[34]

If humans move to a higher altitude, respiration and physical exertion become a problem, but after spending time in high altitude conditions they acclimatize to the reduced partial pressure of oxygen, such as by producing more red blood cells. The ability to acclimatize is an adaptation, but the acclimatization itself is not. Fecundity goes down, but deaths from some tropical diseases also go down. Over a longer period of time, some people are better able to reproduce at high altitudes than others. They contribute more heavily to later generations, and gradually by natural selection the whole population becomes adapted to the new conditions. This has demonstrably occurred, as the observed performance of long-term communities at higher altitude is significantly better than the performance of new arrivals, even when the new arrivals have had time to acclimatize.[35]

Adaptedness and fitness

Fitness-landscape-cartoon
In this sketch of a fitness landscape, a population can evolve by following the arrows to the adaptive peak at point B, and the points A and C are local optima where a population could become trapped.

There is a relationship between adaptedness and the concept of fitness used in population genetics. Differences in fitness between genotypes predict the rate of evolution by natural selection. Natural selection changes the relative frequencies of alternative phenotypes, insofar as they are heritable.[36] However, a phenotype with high adaptedness may not have high fitness. Dobzhansky mentioned the example of the Californian redwood, which is highly adapted, but a relict species in danger of extinction.[25] Elliott Sober commented that adaptation was a retrospective concept since it implied something about the history of a trait, whereas fitness predicts a trait's future.[37]

1. Relative fitness. The average contribution to the next generation by a genotype or a class of genotypes, relative to the contributions of other genotypes in the population.[38] This is also known as Darwinian fitness, selection coefficient, and other terms.
2. Absolute fitness. The absolute contribution to the next generation by a genotype or a class of genotypes. Also known as the Malthusian parameter when applied to the population as a whole.[36][39]
3. Adaptedness. The extent to which a phenotype fits its local ecological niche. Researchers can sometimes test this through a reciprocal transplant.[40]

Sewall Wright proposed that populations occupy adaptive peaks on a fitness landscape. To evolve to another, higher peak, a population would first have to pass through a valley of maladaptive intermediate stages, and might be "trapped" on a peak that is not optimally adapted.[41]

Genetic basis

A large diversity of genome DNAs in a species is the basis for adaptation and differentiation. A large population is needed to carry sufficient diversity. According to the misrepair-accumulation aging theory,[42][43] The misrepair mechanism is important in maintaining a sufficient number of individuals in a species.[44] misrepair is a way of repair for increasing the surviving chance of an organism when it has severe injuries. Without misrepairs, no individual could survive to reproduction age. Thus misrepair mechanism is an essential mechanism for the survival of a species and for maintaining the number of individuals. Although individuals die from aging, genome DNAs are being recopied and transmitted by individuals generation by generation. In addition, the DNA misrepairs in germ cells contribute also to the diversity of genome DNAs.

Types

Adaptation is the heart and soul of evolution.

— Niles Eldredge, Reinventing Darwin: The Great Debate at the High Table of Evolutionary Theory[45]

Changes in habitat

Before Darwin, adaptation was seen as a fixed relationship between an organism and its habitat. It was not appreciated that as the climate changed, so did the habitat; and as the habitat changed, so did the biota. Also, habitats are subject to changes in their biota: for example, invasions of species from other areas. The relative numbers of species in a given habitat are always changing. Change is the rule, though much depends on the speed and degree of the change. When the habitat changes, three main things may happen to a resident population: habitat tracking, genetic change or extinction. In fact, all three things may occur in sequence. Of these three effects only genetic change brings about adaptation. When a habitat changes, the resident population typically moves to more suitable places; this is the typical response of flying insects or oceanic organisms, which have wide (though not unlimited) opportunity for movement.[46] This common response is called habitat tracking. It is one explanation put forward for the periods of apparent stasis in the fossil record (the punctuated equilibrium theory).[47]

Genetic change

Genetic change occurs in a population when natural selection and mutations act on its genetic variability.[48] The first pathways of enzyme-based metabolism may have been parts of purine nucleotide metabolism, with previous metabolic pathways being part of the ancient RNA world. By this means, the population adapts genetically to its circumstances.[10] Genetic changes may result in visible structures, or may adjust physiological activity in a way that suits the habitat.

Habitats and biota do frequently change. Therefore, it follows that the process of adaptation is never finally complete.[49] Over time, it may happen that the environment changes little, and the species comes to fit its surroundings better and better. On the other hand, it may happen that changes in the environment occur relatively rapidly, and then the species becomes less and less well adapted. Seen like this, adaptation is a genetic tracking process, which goes on all the time to some extent, but especially when the population cannot or does not move to another, less hostile area. Given enough genetic change, as well as specific demographic conditions, an adaptation may be enough to bring a population back from the brink of extinction in a process called evolutionary rescue. Adaptation does affect, to some extent, every species in a particular ecosystem.[50][51]

Leigh Van Valen thought that even in a stable environment, competing species constantly had to adapt to maintain their relative standing. This became known as the Red Queen hypothesis, as seen in host-parasite interaction.[52]

Co-adaptation

Plumpollen0060
Pollinating insects are co-adapted with flowering plants.

In coevolution, where the existence of one species is tightly bound up with the life of another species, new or 'improved' adaptations which occur in one species are often followed by the appearance and spread of corresponding features in the other species. These co-adaptational relationships are intrinsically dynamic, and may continue on a trajectory for millions of years, as has occurred in the relationship between flowering plants and pollinating insects.[53][54]

Mimicry

Wasp mimicry
A and B show real wasps; the rest are Batesian mimics: three hoverflies and one beetle.

Bates' work on Amazonian butterflies led him to develop the first scientific account of mimicry, especially the kind of mimicry which bears his name: Batesian mimicry.[55] This is the mimicry by a palatable species of an unpalatable or noxious species, gaining a selective advantage. A common example seen in temperate gardens is the hoverfly, many of which—though bearing no sting—mimic the warning coloration of hymenoptera (wasps and bees). Such mimicry does not need to be perfect to improve the survival of the palatable species.[56]

Bates, Wallace and Fritz Müller believed that Batesian and Müllerian mimicry provided evidence for the action of natural selection, a view which is now standard amongst biologists.[57][58][59]

Trade-offs

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

— Peter Medawar, The Future of Man[60]

All adaptations have a downside: horse legs are great for running on grass, but they can't scratch their backs; mammals' hair helps temperature, but offers a niche for ectoparasites; the only flying penguins do is under water. Adaptations serving different functions may be mutually destructive. Compromise and makeshift occur widely, not perfection. Selection pressures pull in different directions, and the adaptation that results is some kind of compromise.[61]

Since the phenotype as a whole is the target of selection, it is impossible to improve simultaneously all aspects of the phenotype to the same degree.

Consider the antlers of the Irish elk, (often supposed to be far too large; in deer antler size has an allometric relationship to body size). Obviously, antlers serve positively for defence against predators, and to score victories in the annual rut. But they are costly in terms of resource. Their size during the last glacial period presumably depended on the relative gain and loss of reproductive capacity in the population of elks during that time.[63] As another example, camouflage to avoid detection is destroyed when vivid coloration is displayed at mating time. Here the risk to life is counterbalanced by the necessity for reproduction.[64]

Stream-dwelling salamanders, such as Caucasian salamander or Gold-striped salamander have very slender, long bodies, perfectly adapted to life at the banks of fast small rivers and mountain brooks. Elongated body protects their larvae from being washed out by current. However, elongated body increases risk of desiccation and decreases dispersal ability of the salamanders; it also negatively affects their fecundity. As a result, fire salamander, less perfectly adapted to the mountain brook habitats, is in general more successful, have a higher fecundity and broader geographic range.[65]

Pfau imponierend
An Indian peacock's train
in full display

The peacock's ornamental train (grown anew in time for each mating season) is a famous adaptation. It must reduce his maneuverability and flight, and is hugely conspicuous; also, its growth costs food resources. Darwin's explanation of its advantage was in terms of sexual selection: "This depends on the advantage which certain individuals have over other individuals of the same sex and species, in exclusive relation to reproduction."[66] The kind of sexual selection represented by the peacock is called 'mate choice,' with an implication that the process selects the more fit over the less fit, and so has survival value.[67] The recognition of sexual selection was for a long time in abeyance, but has been rehabilitated.[68]

The conflict between the size of the human foetal brain at birth, (which cannot be larger than about 400 cm3, else it will not get through the mother's pelvis) and the size needed for an adult brain (about 1400 cm3), means the brain of a newborn child is quite immature. The most vital things in human life (locomotion, speech) just have to wait while the brain grows and matures. That is the result of the birth compromise. Much of the problem comes from our upright bipedal stance, without which our pelvis could be shaped more suitably for birth. Neanderthals had a similar problem.[69][70][71]

As another example, the long neck of a giraffe is a burden and a blessing. The neck of a giraffe can be up to 2 m (6 ft 7 in) in length.[72] This neck can be used for inter-species competition or for foraging on tall trees where shorter herbivores cannot reach. However, as previously stated, there is always a trade-off. This long neck is heavy and it adds to the body mass of a giraffe, so the giraffe needs an abundance of nutrition to provide for this costly adaptation.[73]

Shifts in function

Adaptation and function are two aspects of one problem.

— Julian Huxley, Evolution: The Modern Synthesis[74]

Pre-adaptation

Pre-adaptation occurs when a population has characteristics which by chance are suited for a set of conditions not previously experienced. For example, the polyploid cordgrass Spartina townsendii is better adapted than either of its parent species to their own habitat of saline marsh and mud-flats.[75] Among domestic animals, the White Leghorn chicken is markedly more resistant to vitamin B1 deficiency than other breeds; on a plentiful diet this makes no difference, but on a restricted diet this preadaptation could be decisive.[76]

Pre-adaptation may arise because a natural population carries a huge quantity of genetic variability.[77] In diploid eukaryotes, this is a consequence of the system of sexual reproduction, where mutant alleles get partially shielded, for example, by genetic dominance.[78] Microorganisms, with their huge populations, also carry a great deal of genetic variability. The first experimental evidence of the pre-adaptive nature of genetic variants in microorganisms was provided by Salvador Luria and Max Delbrück who developed the Fluctuation Test, a method to show the random fluctuation of pre-existing genetic changes that conferred resistance to bacteriophages in Escherichia coli.[79]

Co-option of existing traits: exaptation

Sinosauropteryxfossil
The feathers of Sinosauropteryx, a dinosaur with feathers, were used for insulation, making them an exaptation for flight.

Features that now appear as adaptations sometimes arose by co-option of existing traits, evolved for some other purpose. The classic example is the ear ossicles of mammals, which we know from paleontological and embryological evidence originated in the upper and lower jaws and the hyoid bone of their synapsid ancestors, and further back still were part of the gill arches of early fish.[80][81] The word exaptation was coined to cover these common evolutionary shifts in function.[82] The flight feathers of birds evolved from the much earlier feathers of dinosaurs,[83] which might have been used for insulation or for display.[84][85]

Non-adaptive traits

Some traits do not appear to be adaptive, that is, they have a neutral or deleterious effect on fitness in the current environment. Because genes have pleiotropic effects, not all traits may be functional: they may be what Stephen Jay Gould and Richard Lewontin called spandrels, features brought about by neighbouring adaptations, like the triangular areas under neighbouring arches in architecture which began as functionless features.[86]

Another possibility is that a trait may have been adaptive at some point in an organism's evolutionary history, but a change in habitats caused what used to be an adaptation to become unnecessary or even maladapted. Such adaptations are termed vestigial. Many organisms have vestigial organs, which are the remnants of fully functional structures in their ancestors. As a result of changes in lifestyle the organs became redundant, and are either not functional or reduced in functionality. Since any structure represents some kind of cost to the general economy of the body, an advantage may accrue from their elimination once they are not functional. Examples: wisdom teeth in humans; the loss of pigment and functional eyes in cave fauna; the loss of structure in endoparasites.[87]

Extinction and coextinction

If a population cannot move or change sufficiently to preserve its long-term viability, then obviously, it will become extinct, at least in that locale. The species may or may not survive in other locales. Species extinction occurs when the death rate over the entire species exceeds the birth rate for a long enough period for the species to disappear. It was an observation of Van Valen that groups of species tend to have a characteristic and fairly regular rate of extinction.[88]

Just as there is co-adaptation, there is also coextinction, the loss of a species due to the extinction of another with which it is coadapted, as with the extinction of a parasitic insect following the loss of its host, or when a flowering plant loses its pollinator, or when a food chain is disrupted.[89][90]

Philosophical issues

Adaptation raises philosophical issues concerning how biologists speak of function and purpose, as this carries implications of evolutionary history – that a feature evolved by natural selection for a specific reason – and potentially of supernatural intervention – that features and organisms exist because of a deity's conscious intentions.[91][92] In his biology, Aristotle introduced teleology to describe the adaptedness of organisms, but without accepting the supernatural intention built into Plato's thinking, which Aristotle rejected.[93][94] Modern biologists continue to face the same difficulty.[95][96][97][98][99][100] On the one hand, adaptation is obviously purposeful: natural selection chooses what works and eliminates what does not. On the other hand, biologists want to deny conscious purpose in evolution. The dilemma gave rise to a famous joke by the evolutionary biologist Haldane: "Teleology is like a mistress to a biologist: he cannot live without her but he's unwilling to be seen with her in public.'" David Hull commented that Haldane's mistress "has become a lawfully wedded wife. Biologists no longer feel obligated to apologize for their use of teleological language; they flaunt it."[101]

See also

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Sources

Academy Award for Best Adapted Screenplay

The Academy Award for Best Adapted Screenplay is one of the Academy Awards, the most prominent film awards in the United States. It is awarded each year to the writer of a screenplay adapted from another source (usually a novel, play, short story, TV series, or even another film). All sequels are automatically considered adaptations by this standard (since the sequel must be based on the original story).

See also the Academy Award for Best Original Screenplay, a similar award for screenplays that are not adaptations.

Academy Award for Best Original Score

The Academy Award for Best Original Score is an award presented annually by the Academy of Motion Picture Arts and Sciences (AMPAS) to the best substantial body of music in the form of dramatic underscoring written specifically for the film by the submitting composer.

Adaptation (film)

Adaptation. is a 2002 American comedy-drama metafilm directed by Spike Jonze and written by Charlie Kaufman.The film stars Nicolas Cage as Kaufman and his fictional twin brother Donald, Meryl Streep as Orlean, and Chris Cooper as John Laroche, with Cara Seymour, Brian Cox, Tilda Swinton, Ron Livingston, and Maggie Gyllenhaal in supporting roles.

It is based both on Susan Orlean's non-fiction book The Orchid Thief and Kaufman's experience attempting to adapt the book into a screenplay while suffering from writer's block. Adaptation also adds a number of fictitious elements, including Kaufman's twin brother (also credited as a writer for the film) and a romance between Orlean and Laroche, and culminates in completely invented events including fictional versions of Orlean and Laroche three years after the events related in The Orchid Thief.

Adaptation received awards at the 75th Academy Awards, 60th Golden Globe Awards, and 56th British Academy Film Awards, with Cooper winning the Academy Award for Best Supporting Actor and Kaufman winning the BAFTA Award for Best Adapted Screenplay. The film was ranked in a British Film Institute poll as one of the thirty best films of the 2000s.

Angry Birds

Angry Birds is a video game franchise created by the Finnish company Rovio Entertainment. The series focuses on multi-colored birds which try to save their eggs from green-colored pigs, their enemies. Inspired by Crush the Castle, the game has been praised for its successful combination of fun gameplay, comical style, and low price. Its popularity led to many spin-offs, versions of Angry Birds created for PCs and video game consoles, a market for merchandise featuring its characters, a televised animated series, and a feature film. By January 2014, there were over two billion downloads across all platforms, including both regular and special editions. By July 2015, the series' games were downloaded more than three billion times collectively, making it the most downloaded freemium game series of all time. The original Angry Birds has been called "one of the most mainstream games out right now", "one of the great runaway hits of 2010", and "the largest mobile app success the world has seen so far". An animated feature film based on the series was released by Columbia Pictures on 20 May 2016, and the first main-series sequel, Angry Birds 2, was released on 30 July 2015.

The first game in the series was initially released in December 2009 for iOS. At the time, the swine flu epidemic was in the news, so the staff decided to use pigs as the enemies of the birds. The company released ports of the game to other touchscreen smartphone operating systems, including Android, Symbian, and Windows Phone, and PCs.

Annual plant

An annual plant is a plant that completes its life cycle, from germination to the production of seeds, within one year, and then dies. Summer annuals germinate during spring or early summer and mature by autumn of the same year. Winter annuals germinate during the autumn and mature during the spring or summer of the following calendar year.One seed-to-seed life cycle for an annual can occur in as little as a month in some species, though most last several months. Oilseed rapa can go from seed-to-seed in about five weeks under a bank of fluorescent lamps. This style of growing is often used in classrooms for education. Many desert annuals are therophytes, because their seed-to-seed life cycle is only weeks and they spend most of the year as seeds to survive dry conditions.

Aquatic animal

An aquatic animal is an animal, either vertebrate or invertebrate, which lives in the water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic animals may breathe air or extract oxygen that dissolved in water through specialised organs called gills, or directly through the skin. Natural environments and the animals that live in them can be categorized as aquatic (water) or terrestrial (land). This designation is paraphyletic.

Climate change adaptation

Climate change adaptation is a response to global warming (also known as "climate change" or "anthropogenic climate change"), that seeks to reduce the vulnerability of social and biological systems to relatively sudden change and thus offset the effects of global warming. Even if emissions are stabilized relatively soon, global warming and its effects should last many years, and adaptation would be necessary to the resulting changes in climate. Adaptation is especially important in developing countries since those countries are predicted to bear the brunt of the effects of global warming. That is, the capacity and potential for humans to adapt (called adaptive capacity) is unevenly distributed across different regions and populations, and developing countries generally have less capacity to adapt. Furthermore, the degree of adaptation correlates to the situational focus on environmental issues. Therefore, adaptation requires the situational assessment of sensitivity and vulnerability to environmental impacts.Adaptive capacity is closely linked to social and economic development. The economic costs of adaptation to climate change are likely to cost billions of dollars annually for the next several decades, though the amount of money needed is unknown. Donor countries promised an annual $100 billion by 2020 through the Green Climate Fund for developing countries to adapt to climate change. However, while the fund was set up during COP16 in Cancún, concrete pledges by developed countries have not been forthcoming. The adaptation challenge grows with the magnitude and the rate of climate change.

Another response to climate change is known as climate change mitigation. It advocates to reduce greenhouse gas (GHG) emissions or enhance the removal of these gases from the atmosphere (through carbon sinks). Even the most effective reductions in emissions, however, would not prevent further climate change impacts, making the need for adaptation unavoidable. A study has concluded, with very high confidence, that in the absence of mitigation efforts, the effects of climate change would reach such a magnitude as to make adaptation impossible for some natural ecosystems. Others are concerned that climate adaptation programs might interfere with the existing development programs and thus lead to unintended consequences for vulnerable groups. For human systems, the economic and social costs of unmitigated climate change would be very high.

Color balance

In photography and image processing, color balance is the global adjustment of the intensities of the colors (typically red, green, and blue primary colors). An important goal of this adjustment is to render specific colors – particularly neutral colors – correctly. Hence, the general method is sometimes called gray balance, neutral balance, or white balance. Color balance changes the overall mixture of colors in an image and is used for color correction. Generalized versions of color balance are used to correct colors other than neutrals or to deliberately change them for effect.

Image data acquired by sensors – either film or electronic image sensors – must be transformed from the acquired values to new values that are appropriate for color reproduction or display. Several aspects of the acquisition and display process make such color correction essential – including that the acquisition sensors do not match the sensors in the human eye, that the properties of the display medium must be accounted for, and that the ambient viewing conditions of the acquisition differ from the display viewing conditions.

The color balance operations in popular image editing applications usually operate directly on the red, green, and blue channel pixel values, without respect to any color sensing or reproduction model. In film photography, color balance is typically achieved by using color correction filters over the lights or on the camera lens.

Film adaptation

A film adaptation is the transfer of a work or story, in whole or in part, to a feature film. Although often considered a type of derivative work, recent academic developments by scholars such as Robert Stam conceptualize film adaptation as a dialogic process.

A common form of film adaptation is the use of a novel as the basis of a feature film. Other works adapted into films include non-fiction (including journalism), autobiography, comic books, scriptures, plays, historical sources, and other films. From the earliest days of cinema, in nineteenth-century Europe, adaptation from such diverse resources has been a ubiquitous practice of filmmaking.

Gerard Butler

Gerard James Butler (born 13 November 1969) is a Scottish actor, producer, singer and musician.

After studying law, Butler turned to acting in the mid-1990s with small roles in productions such as Mrs Brown (1997), the James Bond film Tomorrow Never Dies (1997), and Tale of the Mummy (1998). In 2000, he starred as Dracula in the horror film Dracula 2000 with Christopher Plummer and Jonny Lee Miller.

He subsequently played Attila the Hun in the miniseries Attila (2001) and then appeared in the films Reign of Fire with Christian Bale (2002) and Lara Croft: Tomb Raider – The Cradle of Life with Angelina Jolie (2003), before playing André Marek in the adaptation of Michael Crichton's science fiction adventure Timeline (2003). He then was cast as the role of Erik, The Phantom in Joel Schumacher's 2004 film adaptation of the musical The Phantom of the Opera alongside Emmy Rossum. That role earned him a Satellite Award nomination for Best Actor.

Although Attila and The Phantom of the Opera were important breaks, it was only in 2007 that Butler gained worldwide recognition for his portrayal of King Leonidas in Zack Snyder's fantasy war film 300. That role earned him nominations for an Empire Award for Best Actor and a Saturn Award for Best Actor and a win for MTV Movie Award for Best Fight. In the 2010s, he voiced the role of Stoick the Vast in the animated action-fantasy film How to Train Your Dragon, a role he later reprised in Legend of the Boneknapper Dragon (2010), Gift of the Night Fury (2011), How to Train Your Dragon 2 (2014), and How to Train Your Dragon: The Hidden World (2019). He played military leader Tullus Aufidius in the 2011 film Coriolanus, a modernized adaptation of Shakespeare's tragedy of the same name. He also played Sam Childers in the 2011 action biopic Machine Gun Preacher.

Global warming

Global warming is a long-term rise in the average temperature of the Earth's climate system, an aspect of climate change shown by temperature measurements and by multiple effects of the warming. Though earlier geological periods also experienced episodes of warming, the term commonly refers to the observed and continuing increase in average air and ocean temperatures since 1900 caused mainly by emissions of greenhouse gasses in the modern industrial economy. In the modern context the terms global warming and climate change are commonly used interchangeably, but climate change includes both global warming and its effects, such as changes to precipitation and impacts that differ by region. Many of the observed warming changes since the 1950s are unprecedented in the instrumental temperature record, and in historical and paleoclimate proxy records of climate change over thousands to millions of years.In 2013, the Intergovernmental Panel on Climate Change (IPCC) Fifth Assessment Report concluded, "It is extremely likely that human influence has been the dominant cause of the observed warming since the mid-20th century." The largest human influence has been the emission of greenhouse gases such as carbon dioxide, methane, and nitrous oxide. Climate model projections summarized in the report indicated that during the 21st century, the global surface temperature is likely to rise a further 0.3 to 1.7 °C (0.5 to 3.1 °F) to 2.6 to 4.8 °C (4.7 to 8.6 °F) depending on the rate of greenhouse gas emissions and on climate feedback effects. These findings have been recognized by the national science academies of the major industrialized nations and are not disputed by any scientific body of national or international standing.Future climate change effects are expected to include rising sea levels, ocean acidification, regional changes in precipitation, and expansion of deserts in the subtropics. Surface temperature increases are greatest in the Arctic, with the continuing retreat of glaciers, permafrost, and sea ice. Predicted regional precipitation effects include more frequent extreme weather events such as heat waves, droughts, wildfires, heavy rainfall with floods, and heavy snowfall. Effects directly significant to humans are predicted to include the threat to food security from decreasing crop yields, and the abandonment of populated areas due to rising sea levels. Environmental impacts appear likely to include the extinction or relocation of ecosystems as they adapt to climate change, with coral reefs, mountain ecosystems, and Arctic ecosystems most immediately threatened. Because the climate system has a large "inertia" and greenhouse gases will remain in the atmosphere for a long time, climatic changes and their effects will continue to become more pronounced for many centuries even if further increases to greenhouse gases stop.Possible societal responses to global warming include mitigation by emissions reduction, adaptation to its effects, and possible future climate engineering. Almost all countries are parties to the United Nations Framework Convention on Climate Change (UNFCCC), whose ultimate objective is to prevent dangerous anthropogenic climate change. Parties to the UNFCCC have agreed that deep cuts in emissions are required and that global warming should be limited to well below 2.0 °C (3.6 °F) compared to pre-industrial levels, with efforts made to limit warming to 1.5 °C (2.7 °F). Some scientists call into question climate adaptation feasibility, with higher emissions scenarios, or the two degree temperature target.Public reactions to global warming and concern about its effects are also increasing. A 2015 global survey showed that a median of 54% of respondents consider it "a very serious problem", with significant regional differences: Americans and Chinese (whose economies are responsible for the greatest annual CO2 emissions) are among the least concerned.

Internet protocol suite

The Internet protocol suite is the conceptual model and set of communications protocols used in the Internet and similar computer networks. It is commonly known as TCP/IP because the foundational protocols in the suite are the Transmission Control Protocol (TCP) and the Internet Protocol (IP). It is occasionally known as the Department of Defense (DoD) model because the development of the networking method was funded by the United States Department of Defense through DARPA.

The Internet protocol suite provides end-to-end data communication specifying how data should be packetized, addressed, transmitted, routed, and received. This functionality is organized into four abstraction layers, which classify all related protocols according to the scope of networking involved. From lowest to highest, the layers are the link layer, containing communication methods for data that remains within a single network segment (link); the internet layer, providing internetworking between independent networks; the transport layer, handling host-to-host communication; and the application layer, providing process-to-process data exchange for applications.

The technical standards underlying the Internet protocol suite and its constituent protocols are maintained by the Internet Engineering Task Force (IETF). The Internet protocol suite predates the OSI model, a more comprehensive reference framework for general networking systems.

J.C.Staff

J.C.Staff Co., Ltd. (株式会社ジェー・シー・スタッフ, Kabushiki gaisha Jē Shī Sutaffu, J and C stand for "Japan Creative"), is a Japanese animation studio founded in January 1986 by Tomoyuki Miyata, who previously worked at Tatsunoko Production. The studio's first release was Yōtōden in 1987. They have produced several well-known anime series, such as Slayers, Revolutionary Girl Utena, Excel Saga, Alien Nine, Azumanga Daioh, Ikki Tōsen: Battle Vixens, Shingetsutan Tsukihime, Honey and Clover, Shakugan no Shana, The Familiar of Zero, Toradora!, A Certain Magical Index, A Certain Scientific Railgun, Date A Live, Maid Sama!, The Pet Girl of Sakurasou, and Food Wars!: Shokugeki no Soma. Since 2005, the majority of their works have been directed by Takashi Watanabe, Ken'ichi Kasai and Yoshiaki Iwasaki.

List of A Song of Ice and Fire characters

George R. R. Martin's A Song of Ice and Fire novels feature a sizable cast of characters. The series follows three interwoven plotlines: a dynastic war for control of Westeros by several families; the rising threat of the superhuman Others beyond Westeros' northern border; and the ambition of Daenerys Targaryen, the exiled heir of the previous ruling dynasty. The Great Houses of Westeros represent the Seven Kingdoms forged across the continent: the North, the Iron Islands, the Vale of Arryn, the Westerlands, the Stormlands, the Reach, and Dorne. A massive Wall of ice and old magic separates the Seven Kingdoms from the largely unmapped area in the most northern portion of the continent.

Each chapter is narrated in the third-person limited point of view through the eyes of a single character. Beginning with nine POV characters in A Game of Thrones (1996), a total of 31 such characters have narrated over the course of the first five volumes of the series.

One Piece

One Piece (Japanese: ワンピース, Hepburn: Wan Pīsu) is a Japanese manga series written and illustrated by Eiichiro Oda. It has been serialized in Shueisha's Weekly Shōnen Jump magazine since July 22, 1997, and has been collected into 92 tankōbon volumes. The story follows the adventures of Monkey D. Luffy, a boy whose body gained the properties of rubber after unintentionally eating a Devil Fruit. With his crew of pirates, named the Straw Hat Pirates, Luffy explores the Grand Line in search of the world's ultimate treasure known as "One Piece" in order to become the next Pirate King.

The manga spawned a media franchise, having been adapted into an original video animation (OVA) produced by Production I.G in 1998, and an anime series produced by Toei Animation, which began broadcasting in Japan in 1999. Additionally, Toei has developed thirteen animated feature films, one OVA and thirteen television specials. Several companies have developed various types of merchandising and media, such as a trading card game and numerous video games. The manga series was licensed for an English language release in North America and the United Kingdom by Viz Media and in Australia by Madman Entertainment. The anime series was licensed by 4Kids Entertainment for an English-language release in North America in 2004, before the license was dropped and subsequently acquired by Funimation in 2007.

One Piece has received praise for its storytelling, art, characterization, and humor. Several volumes of the manga have broken publishing records, including the highest initial print run of any book in Japan. The official website for Eiichiro Oda's One Piece manga announced that the manga has set a Guinness World Record for "the most copies published for the same comic book series by a single author". As of March 2019, the manga has sold over 450 million copies worldwide, making it the best-selling manga series in history. It became the best-selling manga for the eleventh consecutive year in 2018. One Piece is one of the highest-grossing media franchises of all time, estimated to have generated over $20 billion in total franchise revenue, from the manga, anime, films, games and merchandise.

Pet Sematary (1989 film)

Pet Sematary (sometimes referred to as Stephen King's Pet Sematary) is a 1989 American horror film adaptation of Stephen King's 1983 novel of the same name. Directed by Mary Lambert and written by King, the film features Dale Midkiff as Louis Creed, Denise Crosby as Rachel Creed, Blaze Berdahl as Ellie Creed, Miko Hughes as Gage Creed, and Fred Gwynne as Jud Crandall. Andrew Hubatsek was cast for Zelda's role. King, who scripted from his own book, also has a cameo as a minister. The movie's title is a sensational spelling of "cemetery."

A sequel, Pet Sematary Two, was met with less financial and critical success. A second film adaptation of the same name was released in 2019.

School of Rock

School of Rock is a 2003 comedy film directed by Richard Linklater, produced by Scott Rudin, and written by Mike White. The film stars Jack Black, Joan Cusack, White, and Sarah Silverman. Black plays struggling rock guitarist Dewey Finn, who is kicked out of his band and subsequently disguises himself as a substitute teacher at a prestigious prep school. After witnessing the musical talent of his students, Dewey forms a band of fourth-graders to attempt to win the upcoming Battle of the Bands and pay off his rent.

School of Rock was released on October 3, 2003, by Paramount Pictures, grossing $131 million worldwide on a $35 million budget. The film received positive reviews from critics, with praise for Black's performance. It was the highest grossing music-themed comedy of all time, until it was overtaken in 2015 by Pitch Perfect 2. A stage musical adaptation opened on Broadway in December 2015, and a television adaptation for Nickelodeon premiered on March 12, 2016.

Space adaptation syndrome

Space adaptation syndrome (SAS) or space sickness is a condition experienced by as many as half of all space travelers during their adaptation to weightlessness once in orbit. It is the opposite of terrestrial motion sickness since it occurs when the environment and the person appear visually to be in motion relative to one another even though there is no corresponding sensation of bodily movement originating from the vestibular system.

Stress (biology)

Stress, either physiological or biological, is an organism's response to a stressor such as an environmental condition. Stress is the body's method of reacting to a condition such as a threat, challenge or physical and psychological barrier. Stimuli that alter an organism's environment are responded to by multiple systems in the body. The autonomic nervous system and hypothalamic-pituitary-adrenal (HPA) axis are two major systems that respond to stress.The sympathoadrenal medullary (SAM) axis may activate the fight-or-flight response through the sympathetic nervous system, which dedicates energy to more relevant bodily systems to acute adaptation to stress, while the parasympathetic nervous system returns the body to homeostasis. The second major physiological stress, the HPA axis regulates the release of cortisol, which influences many bodily functions such as metabolic, psychological and immunological functions. The SAM and HPA axes are regulated by several brain regions, including the limbic system, prefrontal cortex, amygdala, hypothalamus, and stria terminalis.Through these mechanisms, stress can alter memory functions, reward, immune function, metabolism and susceptibility to diseases. Definitions of stress differ. One system suggests there are five types of stress labeled "acute time-limited stressors", "brief naturalistic stressors", "stressful event sequences", "chronic stressors", and "distant stressors". An acute time-limited stressor involves a short-term challenge, while a brief natural stressor involves an event that is normal but nevertheless challenging. A stressful event sequence is a stressor that occurs, and then continues to yield stress into the immediate future. A chronic stressor involves exposure to a long-term stressor, and a distant stressor is a stressor that is not immediate.

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