Adaptation

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

Historically, adaptation has been described from the time of the ancient Greek philosophers such as Empedocles and Aristotle. In 18th and 19th century natural theology, adaptation was taken as evidence for the existence of a deity. Charles Darwin proposed instead that it was explained by natural selection.

Adaptation is related to biological fitness, which governs the rate of evolution as measured by change in allele frequencies. Often, two or more species co-adapt and co-evolve as they develop adaptations that interlock with those of the other species, such as with flowering plants and pollinating insects. In mimicry, species evolve to resemble other species; in Müllerian mimicry this is a mutually beneficial co-evolution as each of a group of strongly defended species (such as wasps able to sting) come to advertise their defenses in the same way. Features evolved for one purpose may be co-opted for a different one, as when the insulating feathers of dinosaurs were co-opted for bird flight.

Adaptation is a major topic in the philosophy of biology, as it concerns function and purpose (teleology). Some biologists try to avoid terms which imply purpose in adaptation, not least because it suggests a deity's intentions, but others note that adaptation is necessarily purposeful.

## 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 (a purpose), but thought that it "came about naturally, since such things survived." Aristotle did believe in final causes, but assumed that species were fixed.

In natural theology, adaptation was interpreted as the work of a deity and as evidence for the existence of God. William Paley believed that organisms were perfectly adapted to the lives they led, an argument that shadowed Gottfried Wilhelm Leibniz, who had argued that God had brought about "the best of all possible worlds." Voltaire's satire Dr. Pangloss is a parody of this optimistic idea, and David Hume also argued against design. Charles Darwin broke with the tradition by emphasising the flaws and limitations which occurred in the animal and plant worlds.

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

Other natural historians, such as Buffon, accepted adaptation, and some also accepted evolution, without voicing their opinions as to the mechanism. This illustrates the real merit of Darwin and Alfred Russel Wallace, and secondary figures such as Henry Walter Bates, for putting forward 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.

## General principles

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

### What adaptation is

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

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

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

All adaptations help organisms survive in their ecological niches. The adaptive traits may be structural, behavioural or physiological. Structural adaptations are physical features of an organism, such as shape, body covering, armament, and internal organization. Behavioural adaptations are inherited systems of behaviour, whether inherited in detail as instincts, or as 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.

2\. Adaptedness is the state of being adapted: the degree to which an organism is able to live and reproduce in a given set of habitats.

3\. An adaptive trait is an aspect of the developmental pattern of the organism which enables or enhances the probability of that organism surviving and reproducing.

### What adaptation is not

Adaptation differs from flexibility, acclimatization, and learning, all of which are changes during life which are not inherited. Flexibility deals with the relative capacity of an organism to maintain itself in different habitats: its degree of specialization. Acclimatization describes automatic physiological adjustments during life; learning means improvement in behavioural performance during life.

Flexibility stems from phenotypic plasticity, the ability of an organism with a given genotype (genetic type) to change its phenotype (observable characteristics) in response to changes in its habitat, or to move to a different habitat. The degree of flexibility is inherited, and varies between individuals. A highly specialized animal or plant lives only in a well-defined habitat, eats a specific type of food, and cannot survive if its needs are not met. Many herbivores are like this; extreme examples are koalas which depend on Eucalyptus, and giant pandas which require bamboo. A generalist, on 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 the evolutionary biologist John Maynard Smith.

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

### Adaptedness and fitness

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. However, a phenotype with high adaptedness may not have high fitness. Dobzhansky mentioned the example of the Californian redwood, which is highly adapted, but a relict species in danger of extinction. Elliott Sober commented that adaptation was a retrospective concept since it implied something about the history of a trait, whereas fitness predicts a trait's future.

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. 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.

3\. Adaptedness. The extent to which a phenotype fits its local ecological niche. Researchers can sometimes test this through a reciprocal transplant.

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.

## Types

> Adaptation is the heart and soul of evolution.

### 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. 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).

### Genetic change

Without mutation, the ultimate source of all genetic variation, there would be no genetic changes and no subsequent adaptation through evolution by natural selection. Genetic change occurs in a population when mutation increases or decreases in its initial frequency followed by random genetic drift, migration, recombination or natural selection act on this genetic variation. One example is that the first pathways of enzyme-based metabolism at the very origin of life on Earth may have been co-opted components of the already-existing purine nucleotide metabolism, a metabolic pathway that evolved in an ancient RNA world. The co-option requires new mutations and through natural selection, the population then adapts genetically to its present circumstances. Genetic changes may result in entirely new or gradual change to visible structures, or they may adjust physiological activity in a way that suits the habitat. The varying shapes of the beaks of Darwin's finches, for example, are driven by adaptive mutations in the ALX1 gene. The coat color of different wild mouse species matches their environments, whether black lava or light sand, owing to adaptive mutations in the melanocortin 1 receptor and other melanin pathway genes. Physiological resistance to the heart poisons (cardiac glycosides) that monarch butterflies store in their bodies to protect themselves from predators are driven by adaptive mutations in the target of the poison, the sodium pump, resulting in target site insensitivity. These same adaptive mutations and similar changes at the same amino acid sites were found to evolve in a parallel manner in distantly related insects that feed on the same plants, and even in a bird that feeds on monarchs through convergent evolution, a hallmark of adaptation. Convergence at the gene-level across distantly related species can arise because of evolutionary constraint.

Habitats and biota do frequently change over time and space. Therefore, it follows that the process of adaptation is never fully complete. Over time, it may happen that the environment changes little, and the species comes to fit its surroundings better and better, resulting in stabilizing selection. On the other hand, it may happen that changes in the environment occur suddenly, and then the species becomes less and less well adapted. The only way for it to climb back up that fitness peak is via the introduction of new genetic variation for natural selection to act upon. Seen like this, adaptation is a genetic tracking process, which goes on all the time to some extent, but especially when the population cannot or does not move to another, less hostile area. Given enough genetic change, as well as specific demographic conditions, an adaptation may be enough to bring a population back from the brink of extinction in a process called evolutionary rescue. Adaptation does affect, to some extent, every species in a particular ecosystem.

Leigh Van Valen thought that even in a stable environment, because of antagonistic species interactions and limited resources, a species must constantly had to adapt to maintain its relative standing. This became known as the Red Queen hypothesis, as seen in host-parasite interactions.

Existing genetic variation and mutation were the traditional sources of material on which natural selection could act. In addition, horizontal gene transfer is possible between organisms in different species, using mechanisms as varied as gene cassettes, plasmids, transposons and viruses such as bacteriophages.

### Co-adaptation

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. In other words, each species triggers reciprocal natural selection in the other. These co-adaptational relationships are intrinsically dynamic, and may continue on a trajectory for millions of years, as has occurred in the relationship between flowering plants and pollinating insects.

### Mimicry

Bates' work on Amazonian butterflies led him to develop the first scientific account of mimicry, especially the kind of mimicry which bears his name: Batesian mimicry. This is the mimicry by a palatable species of an unpalatable or noxious species (the model), gaining a selective advantage as predators avoid the model and therefore also the mimic. Mimicry is thus an anti-predator adaptation. A common example seen in temperate gardens is the hoverfly (Syrphidae), many of which—though bearing no sting—mimic the warning coloration of aculeate Hymenoptera (wasps and bees). Such mimicry does not need to be perfect to improve the survival of the palatable species.

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

### 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.

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

> 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 resources. 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. As another example, camouflage to avoid detection is destroyed when vivid coloration is displayed at mating time. Here the risk to life is counterbalanced by the necessity for reproduction.

Stream-dwelling salamanders, such as Caucasian salamander or Gold-striped salamander have very slender, long bodies, perfectly adapted to life at 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.

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." The kind of sexual selection represented by the peacock is called 'mate choice,' with an implication that the process selects the more fit over the less fit, and so has survival value. The recognition of sexual selection was for a long time in abeyance, but has been rehabilitated.

The conflict between the size of the human foetal brain at birth, (which cannot be larger than about 400 cm<sup>3</sup>, else it will not get through the mother's pelvis) and the size needed for an adult brain (about 1400 cm<sup>3</sup>), 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.

As another example, the long neck of a giraffe brings benefits but at a cost. The neck of a giraffe can be up to 2 m (6 ft 7 in) in length. The benefits are that it can be used for inter-species competition or for foraging on tall trees where shorter herbivores cannot reach. The cost is that a long neck is heavy and adds to the animal's body mass, requiring additional energy to build the neck and to carry its weight around.

## Shifts in function

> Adaptation and function are two aspects of one problem.

### 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. Among domestic animals, the White Leghorn chicken is markedly more resistant to vitamin B<sub>1</sub> deficiency than other breeds; on a plentiful diet this makes no difference, but on a restricted diet this preadaptation could be decisive.

Pre-adaptation may arise because a natural population carries a huge quantity of genetic variability. In diploid eukaryotes, this is a consequence of the system of sexual reproduction, where mutant alleles get partially shielded, for example, by genetic dominance. Microorganisms, with their huge populations, also carry a great deal of genetic variability. The first experimental evidence of the pre-adaptive nature of genetic variants in microorganisms was provided by Salvador Luria and Max Delbrück who developed the Fluctuation Test, a method to show the random fluctuation of pre-existing genetic changes that conferred resistance to bacteriophages in Escherichia coli. The word is controversial because it is teleological and the entire concept of natural selection depends on the presence of genetic variation, regardless of the population size of a species in question.

### Co-option of existing traits: exaptation

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. The word exaptation was coined to cover these common evolutionary shifts in function. The flight feathers of birds evolved from the much earlier feathers of dinosaurs, which might have been used for insulation or for display.

## Niche construction

Animals including earthworms, beavers and humans use some of their adaptations to modify their surroundings, so as to maximize their chances of surviving and reproducing. Beavers create dams and lodges, changing the ecosystems of the valleys around them. Earthworms, as Darwin noted, improve the topsoil in which they live by incorporating organic matter. Humans have constructed extensive civilizations with cities in environments as varied as the Arctic and hot deserts. In all three cases, the construction and maintenance of ecological niches helps drive the continued selection of the genes of these animals, in an environment that the animals have modified.

## Non-adaptive traits

Some traits do not appear to be adaptive as they have a neutral or deleterious effect on fitness in the current environment. Because genes often have pleiotropic effects, not all traits may be functional: they may be what Stephen Jay Gould and Richard Lewontin called spandrels, features brought about by neighbouring adaptations, on the analogy with the often highly decorated triangular areas between pairs of arches in architecture, which began as functionless features.

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.

## 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.

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.

## Origin of adaptive capacities

The first stage in the evolution of life on earth is often hypothesized to be the RNA world in which short self-replicating RNA molecules proliferated before the evolution of DNA and proteins. By this hypothesis, life started when RNA chains began to self-replicate, initiating the three mechanisms of Darwinian selection: heritability, variation of type, and competition for resources. The fitness of an RNA replicator (its per capita rate of increase) would likely have been a function of its intrinsic adaptive capacities, determined by its nucleotide sequence, and the availability of resources. The three primary adaptive capacities may have been: (1) replication with moderate fidelity, giving rise to heritability while allowing variation of type, (2) resistance to decay, and (3) acquisition of resources. These adaptive capacities would have been determined by the folded configurations of the RNA replicators resulting from their nucleotide sequences.

## 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. 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. Modern biologists continue to face the same difficulty. On the one hand, adaptation is obviously purposeful: natural selection chooses what works and eliminates what does not. On the other hand, biologists by and large reject 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." Ernst Mayr stated that "adaptedness... is a posteriori result rather than an a priori goal-seeking", meaning that the question of whether something is an adaptation can only be determined after the event.

## See also

- Adaptive evolution in the human genome
- Adaptive memory
- Adaptive mutation
- Adaptive system
- Anti-predator adaptation
- Body reactivity
- Ecological trap
- Evolutionary pressure
- Evolvability
- Intragenomic conflict
- Neutral theory of molecular evolution

Process that fits organisms to their environment


Journey (2012 video game)

Journey is an indie adventure game developed by Thatgamecompany, published by Sony Computer Entertainment, and directed by Jenova Chen. It was released for the PlayStation 3 via PlayStation Network in March 2012 and ported to PlayStation 4 in July 2015. It was later ported to Windows in June 2019 and iOS in August 2019.

In Journey, the player controls a robed figure in a vast desert, traveling towards a mountain in the distance. Other players on the same journey can be discovered, and two players can meet and assist each other, but they cannot communicate via speech or text and cannot see each other's names until after the game's credits. The only form of communication between the two is a musical chime, which transforms dull pieces of cloth found throughout the levels into vibrant red, affecting the game world and allowing the player to progress through the levels. The developers sought to evoke in the player a sense of smallness and wonder and to forge an emotional connection between them and the anonymous players they meet along the way. The music, composed by Austin Wintory, dynamically responds to the player's actions, building a single theme to represent the game's emotional arc throughout the story.

Reviewers of the game praised the visual and auditory art as well as the sense of companionship created by playing with a stranger, calling it a moving and emotional experience, and have since listed it as one of the greatest video games of all time. Journey won several "game of the year" awards and received several other awards and nominations, including a Best Score Soundtrack for Visual Media nomination for the 2013 Grammy Awards. A retail "Collector's Edition", including Journey, Thatgamecompany's two previous titles, and additional media, was released in August 2012.

## Gameplay

In Journey, the player takes the role of a robed figure in a desert. After an introductory sequence, the player is shown the robed figure sitting in the sand, with a large mountain in the distance. The path towards this mountain, the ultimate destination of the game, is subdivided into several sections traveled through linearly. The player can walk in the levels, as well as control the camera, which typically follows behind the figure, either with the analog stick or by tilting the motion-sensitive controller. The player can jump with one button, or emit a wordless shout or musical note with another; the length and volume of the shout depends on how the button is pressed, and the note stays in tune with the background music. These controls are presented pictorially at the beginning of the game; at no point outside of the credits and title screen are any words shown or spoken.

The robed figure wears a trailing magical scarf which allows the player to briefly fly; doing so uses up the scarf's magical charge, represented visually by glowing runes on the scarf. The scarf's runes are recharged by being near floating pieces of red cloth, or a variety of other means. Touching glowing symbols scattered throughout the levels lengthens the initially vestigial scarf, allowing the player to remain airborne longer. Larger strips of cloth are present in the levels and can be transformed from a stiff, dull gray to vibrant red by singing near them. Doing so may have effects on the world such as releasing bits of magic cloth, forming bridges, or levitating the player. This, in turn, allows the player to progress in the level by opening doors or allowing them to reach previously inaccessible areas. The robed figure does not have visible arms to manipulate the game world directly. Along the way, the player encounters flying creatures made of cloth, some of which help the player along. In later levels, the player also encounters hostile creatures made of stone, which upon spotting the player rip off parts of the figure's scarf.

In each level, the player may come across one other player temporarily connected to their game. When players approach each other they charge one another's scarves. They cannot communicate with each other beyond patterns of singing. Players can help each other by activating strips of cloth or showing paths, but cannot hinder each other and are not necessary for completing any level. When two players finish a section at the same time they remain together into the next one; otherwise, they are connected to new players when they move on. While all the figures generally look the same, individual players can be told apart by unique symbols which are shown floating in the air when they sing and are displayed on their robes at all times. Players may also gain decorative patterns on their robe with successive playthroughs which can be distinguishing. The entire game takes two to three hours to complete.

## Story

Journey is a wordless story told through gameplay and visual-only cutscenes. The player's character begins near a small sand dune in a vast desert. Walking to the top of the dune, the character can see looming in the far distance a large mysterious mountain with a glowing crevice that splits its peak. As the character approaches the mountain, they find the remnants of a once-thriving civilization, eroded by sand over time. Scattered throughout the ruins at the end of each area are stones where the traveler rests and has visions of meeting a large, white-robed figure in a circular room. Art adorns the walls, describing the rise and fall of the player character's civilization, which also mirrors the player's journey. As the player journeys into the remains of a once sprawling city at the base of the mountain, they find they must also contend with roaming, ancient, and hostile automaton weapons left over from a war that ended the civilization.

A vision shows the traveler crumble before reaching their destination, but the traveler chooses to continue on. Eventually making it safely to the mountain itself, the traveler begins to make their way up it, struggling as they enter the colder climates and encounter deep snow and high winds. With the crevice still a fair distance away, the traveler falls and collapses in the snow. Six of the white-robed figures appear before the character and grant the traveler new energy, allowing the player to reach the summit of the mountain and walk into the crevice as the screen fills with white. The player is then shown the game's credits, playing over the ending cinematic scene. This scene shows a shooting star emanating from the crevice and traversing the path the traveler took through the ruins, and shows glimpses of other robed travelers heading towards the mountain. Eventually, the star comes to rest at the sand dune where the game began, and the player is given the option of starting the game again. As the credits end, the player is shown the usernames of the other travelers who shared part of the journey.

## Development

Journey was the last game made under a three-game contract between Thatgamecompany and Sony Computer Entertainment, the first two being Flow and Flower. Development of the game began in 2009, after the release of Flower. The 18-person development team for Journey was composed mainly of creators of the company's previous games; co-founder Jenova Chen was the creative director and Nick Clark returned as lead designer. Kellee Santiago, producer of Flow and Flower, did not reprise her duties, concentrating instead on her role as the company's president, and was replaced by Robin Hunicke.

When development began, Sony expected the game to be completed in a year, rather than the more than three it finally took. Thatgamecompany always expected to need an extension; according to Hunicke, they believed finishing the game within a year was "unrealistic". Development ended up taking even longer than anticipated, as the team had difficulties paring down their ideas for the game and maintaining efficient communication. Over the course of development the team grew from seven to eighteen people. At the end of the second year, when Sony's extension had run out, the game did not spark the emotions in the player that the team wanted. Sony agreed to another one-year extension, but development ultimately exceeded even that.

The stress of the project led to the feeling there was not enough time or money to complete everything the team wished to, which added to the stress and caused arguments about the design of the game. The developers ended up reducing the overtime they spent on the project to avoid burning out, though it meant further delays and risked the company running out of money as the game neared completion. In a speech at the 16th annual D.I.C.E. Awards in 2013, Chen admitted the company had indeed been driven to bankruptcy in the final months of development and that some of the developers had gone unpaid at the time. Hunicke described the solution to finally finishing the game as learning to let go of tensions and ideas that could not make it into the game and be "nice to each other".

The game is intended to make the player feel "small" and to give them a sense of awe about their surroundings. The basic idea, as designed by Chen, was to create something that moved beyond the "typical defeat/kill/win mentality" of most video games. The team initially created a prototype named Dragon that involved players trying to draw away a large monster from other players but eventually discarded it after finding it was too easy for players to ignore each other in favor of their own objectives.

The developers designed Journey like a Japanese garden, where they attempted to remove all the elements that did not fit, so the emotions they wanted to evoke would come through. This minimalism is intended to make the game feel intuitive to the player, so they can explore and feel a sense of wonder without direct instructions. The story arc is designed to explicitly follow Joseph Campbell's monomyth theory of narrative, or hero's journey, as well as to represent the stages of life, so as to enhance the emotional connection of the players as they journey together. In his D.I.C.E. speech, Chen noted that 3 of their 25 testers had cried upon completing the game.

The game's desert setting is largely based on the Middle East, and incorporates Arabic culture, art and architecture. Jenova Chen and art director Matt Nava did not want the setting to be too Western or Eastern, so they felt the Middle East was an ideal middle ground. In addition, Journey also incorporates Chinese and Tibetan cultural influences, drawing from Chen's childhood in China.

The multiplayer component of Journey was designed to facilitate cooperation between players without forcing it, and without allowing competition. It is intended to allow the players to feel a connection to other people through exploring with them, rather than talking to them or fighting them. The plan was "to create a game where people felt they are connected with each other, to show the positive side of humanity in them". The developers felt the focus on caring about the other player would be diluted by too many game elements, such as additional goals or tasks, as players would focus on those and "ignore" the other player. They also felt having text or voice communication or showing usernames would allow players' biases and preconceptions to come between them and the other player.

Journey was released on March 13, 2012, for download on the PlayStation Network. A PlayStation Home Game Space, or themed area, based on Journey was released on March 14, 2012, and is similar in appearance to the game. A retail "Collector's Edition" of the game was released on August 28, 2012. In addition to Journey, the disc-based title includes Flow and Flower; creator commentaries, art, galleries, and soundtracks for all three games; non-related minigames; and additional content for the PlayStation 3. In September 2012, Sony and Thatgamecompany released a hardcover book entitled "The Art of Journey", by the game's art director Matt Nava, containing pieces of art from the game ranging from concept art to final game graphics.

On July 21, 2015, Journey was released on the PlayStation Network for the PlayStation 4, ported by United Kingdom studio Tricky Pixels; owners of the digital PlayStation 3 version of the game were able to download the new version for free. The PlayStation 4 version of the game features improved graphics over the original, with a higher resolution and frame rate and improved texture quality. According to Tricky Pixels, the original PlayStation 3 game was "a masterpiece of PlayStation 3 programming" and porting the game to the PlayStation 4 was "an immense technical challenge". Annapurna Interactive published ports of Journey for Windows and iOS in 2019, developed by Inline Assembly.

### Music

The music in Journey was composed and orchestrated by Austin Wintory, who had previously worked with Thatgamecompany on the soundtrack for Flow. Wintory worked closely on the soundtrack with sound designer Steve Johnson, as well as the programming team, so the music would dynamically tie in to both the actions of the player and to sound effects caused by nearby game objects, and feel as if it were "unfolding in real time". Johnson felt having short pieces of music that looped without reacting to the player would be a "missed opportunity", and wanted to create music that changed while still containing a composed emotional arc. Jenova Chen met with Wintory at the start of the game's development to describe his vision for the project, and Wintory left the meeting and created the core of the main theme before he reached his car, and composed and recorded the main cello theme for the soundtrack that night. He continued to work on the soundtrack for the next three years in a collaboration with the development team; he would create a track, which the team would use while creating an area in the game, and Wintory would play the section while revising the music and then send it back. Wintory spent time experimenting and discarding many ideas; while the first track, "Nascence", came easily, the final track, "Apotheosis", went through several widely varied attempts.

Unlike many games, where different songs have different themes for each character or area, Wintory chose to base all the pieces on one theme which stood for the player and their journey, with cello solos especially representing the player. Wintory describes the music as "like a big cello concerto where you are the soloist and all the rest of the instruments represent the world around you", though he describes it as not necessarily orchestral due to the inclusion of electronic aspects. The cello begins the game as "immersed in a sea of electronic sound", before first emerging on its own and then merging into a full orchestra, mirroring the player's journey to the mountain. Whenever the player meets another person, harps and viola are dynamically incorporated into the music. While the game's art style is based on several different cultures, Wintory tried to remove any overt cultural influences from the music to make it "as universal and culture-less as possible". Tina Guo features as the cellist for the soundtrack. She is a close friend of Wintory and has performed "Woven Variations" with him, an eight-minute live orchestral variation on the Journey soundtrack. All the non-electronic instruments in the soundtrack were recorded with the Macedonia Radio Symphonic Orchestra in Skopje, North Macedonia. A "Woven Variations" performance influenced the ending of the game: at the conclusion of development, Wintory was having difficulty with the ending to "Apotheosis", the final track of the game, while the development team was unsure how to end the player's journey at the top of the mountain. While they were planning a large, dramatic conclusion to both, in the concert Wintory had the orchestra fall away at the end of the piece to showcase Guo's cello performance. Inspired, Wintory and the team ended "Apotheosis" and the game the same way, with the game world fading away to leave only the player.

The soundtrack was released as an album on April 10 on iTunes and the PlayStation Network. The album is a collection of the soundtrack's "most important" pieces, arranged by Wintory to stand alone without the context of the player's actions. The album comprises 18 tracks and is over 58 minutes long. It features the voice of Lisbeth Scott for the final track, "I Was Born for This". After its release, the soundtrack reached the top 10 of the iTunes Soundtrack charts in more than 20 countries. It also reached No. 116 on the Billboard sales charts, with over 4000 units sold in its first week after release, the second-highest position of any video game music album to date. The soundtrack was released as a physical album by Sumthing Else Music Works on October 9, 2012. In 2012 Wintory released a download-only album of music on Bandcamp titled Journey Bonus Bundle, which includes variations on themes from Journey and Flow. The soundtrack itself was subsequently released on Bandcamp on June 19, 2013. An album of piano arrangements titled Transfiguration was released on May 1, 2014, on Bandcamp as both a digital and a physical album. A two-record vinyl version of the album was released in 2015.

In January 2016, Wintory started a Kickstarter for a Journey Live concert tour, in which the fifteen-piece Fifth House Ensemble from Chicago will perform the music from the game while a player works their way through the game. The ensemble will react to the player's actions, using a specially-scored version of the soundtrack, composed by Patrick O'Malley with Wintory's oversight, that breaks the music into small pieces to enable this reaction. Wintory had wanted to do a performance of the Journey soundtrack in this interactive manner but did not have the time to rework the soundtrack for this purpose. Wintory came to know Dan Visconti, the composer for Fifth House Ensemble, after Visconti published his praise for the Journey soundtrack and had encouraged other members of the ensemble to play the game. The group saw how Journey's soundtrack had been used for various Video Games Live concerts and believed they could pull off Wintory's vision of an interactive concert, doing most of the reworking of the soundtrack under Wintory's direction. Sony provided Wintory with a modified version of the game with the music disabled for the concert performance. The Kickstarter was launched for \$9,000 in funding for a four-city tour, but within a few days already surpassed its funding levels, allowing for more cities to be included.

In March 2022, on the game's tenth anniversary, Wintory released a re-arranged version of Journey's soundtrack for a full orchestra performed by the London Symphony Orchestra, the London Voice choir, and cellist Tina Guo. Titled Traveller: A Journey Symphony, Wintory re-imagined elements of the soundtrack to use a larger set of instruments and players and a greater number of vocalists, taking the number of performers from around 30 on the original soundtrack to around 130, while trying to stay true to the original compositions. The album was funded and co-produced by Wintory and Andrea Pessino, co-founder of video game developer Ready at Dawn, for whose games Wintory has composed. It was released as a digital album and on vinyl.

## Reception

Journey received critical and commercial success worldwide. After its release, it became the fastest-selling game to date on PlayStation Store in both North America and Europe. At E3 2011, prior to release, the game won awards for best downloadable game from 1UP.com, GameSpy, and GameTrailers. After publication, the game was heavily honored at end of the year awards. Journey was selected as the best game of the year by IGN and GameSpot, among others. Journey was an Interactive Narrative and Game + Play 2021 Peabody Award Winner.

The soundtrack was nominated for the Best Score Soundtrack for Visual Media at the 2013 Grammy Awards, the first video game soundtrack to be nominated for that category, though it did not win. Additionally, the game won the award for best music and was nominated for the best graphics award from IGN, and was selected as the best PlayStation Network game by GameSpot. At the Spike Video Game Awards, Journey won awards as the best PlayStation 3 game, the best indie game, and the game with the best music, and was additionally nominated for game of the year, best downloadable game, best graphics, and best song in a game for "I Was Born For This". It received the 2013 Annie Award for video game animation. It won five awards at the 2013 British Academy of Film and Television Arts awards: Artistic Achievement, Audio Achievement, Game Design, Online Multiplayer, and Original Music, and was nominated for Best Game, Game Innovation and Story. In March 2013, it won six awards at the annual Game Developers Choice Awards: Best Audio, Best Game Design, Best Visual Arts, Best Downloadable Game, the Innovation Award, and Game of the Year. At the 16th Annual D.I.C.E. Awards, Journey won eight awards, the most honors received of the night, which includes "Game of the Year", "Outstanding Innovation in Gaming", "Casual Game of the Year", "Outstanding Achievement in Game Direction", "Outstanding Achievement in Art Direction", "Outstanding Achievement in Online Gameplay", "Outstanding Achievement in Original Music Composition", and "Outstanding Achievement in Sound Design"; it was additionally nominated for "Downloadable Game of the Year", "Outstanding Achievement in Gameplay Engineering", and "Outstanding Achievement in Story".

Journey received high acclaim from critics who praised the visual and auditory art direction as well as the emotional response playing with a stranger created. It received the IGN Best Overall Game Award for 2012 and Ryan Clements of IGN described the game as "the most beautiful game of its time", saying, "each moment is like a painting, expertly framed and lit". Jane Douglas of GameSpot concurred, calling it "relentlessly beautiful" and lauding the visual diversity of the world and the depiction of the rippling sand; Matt Miller of Game Informer added praise for the animation of the sand and creatures, saying the game was visually stunning. The music was also complimented, with Miller describing it as a "breathtaking musical score" and Douglas calling it "moving, dynamic music".

Reviewers were especially pleased with the emotional experience of playing the game, particularly with other players. Christian Donlan of Eurogamer described it as a "non-denominational religious experience" that, with the addition of another player, moves beyond metaphors and becomes a "pilgrimage" to the player. A reviewer writing for Edge magazine said the emotional arc of the game hits with "occasionally startling power", while Patrick Shaw from Wired said the game made him feel a "wide range of emotions ... wonder, fear, even sadness". Miller said all three times he played the game, "each time, without fail, individual moments... managed to give me goosebumps, and those moments have remained on my mind for weeks afterward". Joel Gregory of PlayStation Official Magazine praised the game's story for being open to the player's interpretation, leaving an ambiguity that drew him in. The addition of an unnamed second player was described by Donlan as brilliant and as a "master stroke", and Edge said it made for "a more absorbing, more atmospheric experience". In 2019, Journey was ranked 48th on The Guardian newspaper's The 50 Best Video Games of the 21st Century list.

The few criticisms for the game centered on its length and pacing. Clements noted that not all players would appreciate a game with a "deliberate, melancholic pace" and short duration, comments echoed by the Edge review. Miller noted the lack of complex gameplay elements in Journey, and Shaw was disappointed that the game was only a few hours long, though Douglas said the length was perfect. Miller concluded the game could be compared to "a musical concert, a well-directed film, or a long-awaited book", while Clements concluded, "completing Journey will create memories that last for years".

2012 video game

Annie Award for Best Animated Video Game winners

British Academy Games Award for Game Design winners

British Academy Games Award for Multiplayer winners

D.I.C.E. Award for Game of the Year winners

D.I.C.E. Award for Online Game of the Year winners

D.I.C.E. Award for Outstanding Achievement in Art Direction winners

D.I.C.E. Award for Outstanding Achievement in Audio Design winners

D.I.C.E. Award for Outstanding Achievement in Game Direction winners

D.I.C.E. Award for Outstanding Achievement in Original Music Composition winners

Game Developers Choice Award for Game of the Year winners

Multiplayer and single-player video games

Video games developed in the United States


Jurassic World Evolution

Jurassic World Evolution is a construction and management simulation video game developed and published by Frontier Developments. Based on the 2015 film Jurassic World, the game was released in June 2018, for PlayStation 4, Windows, and Xbox One. A Nintendo Switch port of the game was released in November 2020. In the game, players construct a dinosaur park on Las Cinco Muertes Archipelago, a group of five islands also known as the "Five Deaths". The game features more than 40 types of dinosaurs; their genes can be modified to introduce new features. Players are given contracts to fulfill by three divisions, Science, Security and Entertainment, allowing them to progress. A sandbox mode set on Isla Nublar, the setting of the first and fourth films, can be unlocked. It can also be used from the main menu without having to be unlocked.

The game was created by a development team of approximately 100 people with a budget of around £8 million. Its development began in 2016, after NBCUniversal approached Frontier Developments about creating a game to accompany the theatrical release of Jurassic World: Fallen Kingdom. To do this, Frontier staff inspected different dinosaur models and reference materials sent by Universal, watched the Jurassic Park films, and read the novels and fan theories. The team consulted paleontologist Jack Horner when they designed the dinosaurs. Jeff Goldblum, Bryce Dallas Howard and BD Wong reprised their roles from the Jurassic Park film series, voicing remarks to players and contributing to the game's narrative.

Announced at Gamescom 2017, the game received a generally mixed critical reception. Critics praised its dinosaur designs and graphics, but the game's contracts, simulation and management gameplay were less well received. The game's tutorial and learning curve were also criticized. Seven months after its initial release, the game had sold two million copies through digital and physical sales, making it the most successful game launched by Frontier. The game was supported with free updates and downloadable content upon release. As of March 2020, the game had sold three million copies. A sequel, Jurassic World Evolution 2, was released on November 9, 2021.

## Gameplay

Jurassic World Evolution is a business simulation game that allows the player to construct a Jurassic World dinosaur theme park with attractions and research facilities. Players must build an Expedition Center, which sends paleontologists to fossil dig sites to obtain dinosaurs' DNA material. DNA sequencing, which can be done in the Fossil Center unlocks new dinosaurs and updates their statistics, such as lifespan and resilience. With enough DNA content, players can use the Hammond Creation Lab to breed and incubate dinosaurs. Players can also improve the dinosaurs' genes by integrating DNA from modern species with that of the dinosaurs to fill their gaps and allow them to evolve. Modifications to the dinosaurs' DNA change their base statistics, as well as everything from their level of aggressiveness to their appearance. The game features a terrain tool which allows players to modify the environment by planting trees and creating water sources.

Dinosaurs are the game's main attraction and income earner, although money can also be earned through selling products in stores and guests staying at hotels. The game features approximately 40 dinosaur species at launch. Players can name each dinosaur after they are incubated. Players need to build enclosures to contain dinosaurs for visitors' viewing. The needs of different dinosaurs, like the type of food they eat and the extent of the social interactions they require, must be met to keep them healthy and satisfied. Dinosaurs, controlled by artificial intelligence, will interact with each other and the environment, as well as guests if they have broken out of their enclosure. For instance, carnivores will attack carnivores of a different species, and they will hunt down herbivores. Players also need to construct various entertainment rides, as well as amenities like restaurants and shops to please the guests. An example of tourist attractions is the Gyrosphere or the monorail from Jurassic World. Players can also use the game's photo mode to take pictures of dinosaurs, which help the park to earn money and publicity. Each entertainment facility and amenity comes with its own management system. Players are able to set and adjust entry fees as well as the number of staff present in each facility. Dinosaurs can be sold to earn additional income.

Various emergency situations may happen in the park, including power failures, unpredictable weather, and dinosaur breakouts, which must be addressed by players to ensure guests' safety and happiness. Players can build an ACU Center and a Ranger Station, which are responsible for maintaining the park's security. They can sedate escaped dinosaurs, medicate sick dinosaurs, resupply dinosaur feeders, transport dinosaurs, fix fences, and more. Players can also control vehicles from a third-person perspective such as helicopters and 4x4 trucks to complete these tasks. Emergency shelters to protect the guests, as well as other security structures like power network redundancies and storm warning centers, can be built. Many of these security facilities can be upgraded to strengthen their efficiency when dealing with emergencies.

### Gameplay modes

In the career mode, the player's goal is to develop five-star parks across the fictional islands of the Las Cinco Muertes Archipelago. Various characters from the Jurassic World franchise assist players throughout the game. Players will meet other key figures representing the three branches of the park's development: Entertainment, Security, and Science. Each of these characters tries to convince players to develop the park in accordance with their advice. They give players "contracts" to complete which include a series of goals and objectives. These contracts add narrative to the game, as well as provide rewards and reputation in their respective domains. Players are advised to keep a close eye on their reputation within each division. If a player's reputation within a division gets too low, that creates a sabotage in the player's park that will need to be attended to immediately. For instance, the park's power could be shut down allowing dinosaurs to break out, or a disease could be introduced to infect them. These divisions all feed into the parks' ratings. The five islands, each with different characteristics and challenges, will gradually unlock with sufficient positive park ratings.

Isla Nublar — the island featured in Jurassic Park, Jurassic World, and Jurassic World: Fallen Kingdom — is the setting for a sandbox mode that is separate from the game's career mode. The sandbox mode is unlocked once a four-star park rating is achieved on Isla Matanceros, the starting island. Once this is accomplished, everything players have unlocked in career mode, such as building upgrades and dinosaurs, will transfer over to the sandbox; anything locked in career mode remains locked in the sandbox. In sandbox mode, players have unlimited funds, and they can set the weather and time of day at their parks. Challenge Mode, available in an update after the game's release, involves playing with adjustable levels of difficulty and limited money, in addition to other differences like fees and penalties against players.

There is a mode available from the main menu, which allows every island to be played in sandbox mode.

## Development and release

Jurassic World Evolution was developed and published by Frontier Developments, and is based on the 2015 film Jurassic World, although the game is not considered canon, which allowed for more creative freedom. NBCUniversal had wanted a game to accompany the theatrical release of its 2018 film Jurassic World: Fallen Kingdom and approached Frontier Developments about creating it two years or so before the film's eventual release date. Frontier Developments was also interested in creating a dinosaur game.

The game was created with a development team of approximately 100 people, on a budget of around £8 million. It was built using Frontier Developments' Cobra game engine. Universal Pictures provided the developers with dinosaur models from the films to allow for a high level of detail, as well as reference materials and audio. To aid in creating the game, the development team closely watched the films and read Michael Crichton's novels, Jurassic Park and The Lost World, and fan theories. In addition, Universal and the team had discussions about various elements in the game. History from each of the films was added to the game. For example, Jeff Goldblum reprised his role as Ian Malcolm from the first two Jurassic Park films. Bryce Dallas Howard and BD Wong also reprised their roles from Jurassic World.

Tyrannosaurus was one of the first dinosaurs the development team began working on. For the T. rex roars, the game's sound team obtained audio samples from the films and then altered them for originality. The animation team then altered their T. rex roaring animations to fit the sound effects. To design the dinosaurs the development team primarily referred to the films for consistency, while incorporating some of the latest scientific discoveries to add to their designs. The team also studied birds and other animals to aid in designing the dinosaurs. In addition to their designs, dinosaur behavior was based on a combination of the films and scientific findings. Paleontologist Jack Horner, who served as an advisor on the films, was also consulted for advice on the game's dinosaurs. They were given bright and colorful hides based on new dinosaur research.

In February 2017, Frontier's CEO David Braben announced the company was working on a new project based on an existing Hollywood property. The game was announced during Gamescom 2017 held in August. Later, on 7 October, footage showcasing the in-game engine was revealed during the first annual Frontier Expo. The game was digitally released for Microsoft Windows, PlayStation 4 and Xbox One on 12 June 2018, coinciding with the theatrical release of Jurassic World: Fallen Kingdom. Physical copies of the PlayStation 4 and Xbox One version were distributed by Sold Out beginning on 3 July.

### Updates

Several updates were released for the game throughout 2018, and Frontier collaborated with Universal on each of them. A free game update based on Jurassic World: Fallen Kingdom, with six dinosaurs from the film, was released on 22 June. In August, Frontier Developments announced an upcoming patch that would alter the sizes of several dinosaurs to match their real-life and film counterparts. The update was released in September, and included additional sandbox and gameplay options, as well as the addition of Challenge Mode.

The first paid downloadable content (DLC), Secrets of Dr. Wu, was released on 20 November. The DLC introduced new story missions, research options and new dinosaur and hybrid species. On the same day, Frontier introduced new AI behaviors and a day-night cycle into the game via a free update. Frontier released the Cretaceous Dinosaur Pack and the Carnivore Dinosaur Pack in December 2018 and April 2019 respectively. Each dinosaur pack introduces three new dinosaurs species. In 2019, a paid DLC titled Claire's Sanctuary was released on 18 June. Set after Fallen Kingdom, the expansion features a standalone campaign which sees players relocating the remaining dinosaurs trapped on Isla Nublar to Sanctuary Island.

Another paid DLC pack, titled Return to Jurassic Park, was released on 10 December. The DLC includes park features and locations from the original Jurassic Park featured in the first film. The DLC also includes Isla Sorna, the island featured in the films The Lost World: Jurassic Park (1997) and Jurassic Park III (2001). It also features seven new missions with new voice work by Goldblum, as well as Sam Neill and Laura Dern, the latter reprising their roles as Dr. Alan Grant and Dr. Ellie Sattler. The DLC reunites the three actors for the first time since the original 1993 film. Return to Jurassic Park features an original story that takes place shortly after the first film, ignoring the sequels. In the story, Grant, Malcolm and Sattler return to Isla Nublar and attempt to get the park operational.

For months, Frontier tried to arrange for Neill, Goldblum and Dern to record their lines together, but scheduling issues prevented this from occurring. Instead, recordings of one actor would be played for the others to aid them in recording their own lines. The DLC also features the character of John Hammond, who was portrayed in the films by Richard Attenborough (1923–2014). In the DLC, Hammond was portrayed by voice actor Mackenzie Gray. Return to Jurassic Park was in development for a while as the Frontier team wanted to take its time to create a comprehensive DLC based on the first film. It is the biggest update created for Jurassic World Evolution. Executive producer Rich Newbold said: "Essentially, the whole game has been rebuilt with a Jurassic Park version of every building". Dinosaur designs based on The Lost World: Jurassic Park and Jurassic Park III were also introduced in the DLC. For reference, Universal provided the developers with original assets and audio files from the film production archives to ensure that the dinosaurs move, sound, and look like their film counterparts. The package introduced Compsognathus and Pteranodon, as well as new designs for the game's Tyrannosaurus rex and Velociraptor, matching their original appearance in the first film.

A Nintendo Switch port of the game, titled Jurassic World Evolution: Complete Edition, was released on 3 November 2020. It includes all the DLC packs and updates that had been previously released for the other platforms.

## Reception

Critics praised the dinosaurs featured in the game. Sam Loveridge of GamesRadar+ liked the variety as well as the cutscene that was displayed when a dinosaur is released from the incubation center. She enjoyed being able to control the jeep and the ACU helicopter from a third-person perspective, a mechanic she applauds for allowing players to relate to the dinosaurs. She added that she "lost entire evenings to [the game] without even thinking" due to its relaxing nature. Game Revolution's Paul Tamburro praised the dinosaurs' design, in particular, Frontier's attention to detail and the dinosaurs' animation. James Swinbanks of GameSpot agreed, praising the dinosaurs. He also enjoyed the need to learn each dinosaur's personality, requirements and behaviors, adding the process is "surprisingly satisfying". Game Informer 's Daniel Tack liked that players can experiment with different genes, though he commented that it was not a "freeform experience". Dan Stapleton of IGN disagreed and felt that unlocking genes was tedious, describing the process as a series of "mandatory robotic actions". While he liked the dinosaur variety, he lamented the lack of pterosaurs and aquatic prehistoric reptiles at launch. Destructoid's Dan Roemer commended the inclusion of lesser- known dinosaurs species, singling out Giganotosaurus and Deinonychus.

The simulation aspect of the game received mixed reviews. Loveridge held a positive opinion regarding dinosaur management, as each species has their own needs and niches that players must fulfill. She felt the park management aspect, including the construction of facilities and utilities, was "minimal". Tamburro compared the simulation to RollerCoaster Tycoon and Zoo Tycoon. He enjoyed the chaos created when dinosaurs break out, but he was disappointed that dinosaurs did not attack the park's staff carrying out maintenance inside the enclosures. Philippa Warr of PC Gamer praised the dinosaurs but criticized the lack of attention to missions and individual guests' views and happiness ratings on the park. She also noted the simulation lacked depth, as buildings looked largely the same and the environment soon became stale. Swinbanks felt dealing with natural disasters and dinosaur breakout was exciting initially, but soon became repetitive. He also lamented the lack of new challenges presented in the later stage of the game. Tack criticized the tiresome aspect of simulation and the constant need to wait for objectives to be completed, comparing them to a mobile game. Describing the simulation as "shallow", Roemer criticized the lack of time constraints, cleanliness ratings, and a full day-night cycle. He was also disappointed by the small size of each island, which made park building "unfulfilling". Stapleton agreed, saying that the small islands limited players' creativity. He noted that players cannot speed up time in the game, a signature feature in many other simulators and builders, and criticized its exclusion as it forced players to wait aimlessly to get enough cash to perform an action.

The game's contracts garnered mixed opinions. Loveridge believed they helped introduce a structure to the game, though she commented that these missions did not form a cohesive narrative. Tamburro enjoyed the process of progressing from one island to the next, as each island has its own unique layout and landscape prompting players to create a new park that is different from the previous ones. He noted that the contracts helped players to unlock new items, but he felt they were not "exciting". Warr criticized the lack of variety featured in the missions, with different scenarios only presenting minor modifications and adjusting the difficult slightly. She also disliked the contracts for occasionally forcing players to complete missions that do not make sense, such as releasing a dinosaur to kill guests. Swinbanks also criticized three contract factions, adding that players' need to satisfy and balance all three parties demands as "arbitrary". Roemer praised Goldblum's performance, though he noted the overarching plot "goes nowhere". He described the game's progression system as "awful" and wished to skip them entirely.

The game's tutorial and guidance to players was criticized. Loveridge noted that certain missions were confusing with some late game objectives being presented too early, forcing him to dismiss them to progress. She singled out the tutorial section for being incompetent, failing to inform players about key aspects of the game such as power distribution and landscaping tools, which often create obstruction. and terrain errors that prevent players from constructing certain buildings and modifying the landscape. Warr noted pacing errors with the tutorials, saying that some of them showed up way too late.

Reviewers had mixed opinions of the game as a whole. Loveridge called it an intricate simulation game that fans of the series would enjoy, and she commended the many references to the film featured in the game. Tamburro agreed and called it the best Jurassic Park-themed game, though he noted that the game had more constraints than Frontier's previous park builder, Planet Coaster. Initially impressed by the game, Warr was disappointed after extended play time feeling the game was lacking depth. Swinbanks noted that despite its shortcomings, the game was "faithful" to the franchise. He felt that it was "a good park management sim in its own right". Roemer felt that the game's development was rushed, and the lack of depth was problematic. Stapleton called it a "bad" game for being largely boring.

### Sales

Five weeks after its initial release, the game had sold one million copies through digital and physical sales. Seven months after the game's initial release, Frontier declared the game its biggest launch and revealed that more than 2 million copies were sold. As of March 2020, three million copies of the game had been sold.

### Accolades

The game was nominated for "Best Audio Design" at the 2018 Golden Joystick Awards, losing to God of War. It won the award for "Game, Simulation" at the National Academy of Video Game Trade Reviewers Awards, and was nominated for "Excellence in Convergence" at the SXSW Gaming Awards, losing to Marvel's Spider-Man. It was also nominated for "Best Game Design" and "Best Audio" at the Develop:Star Awards.

## Sequel

A sequel, Jurassic World Evolution 2, was announced by Frontier Developments in June 2021, and released on November 9. Goldblum reprises his role for the game, which includes additional features, pterosaurs, and marine reptiles. The game takes place after the events of Jurassic World: Fallen Kingdom, and was released for PC, PlayStation 4, PlayStation 5, Xbox One, and Xbox Series X and Series S.

## See also

- Jurassic Park III: Park Builder
- Jurassic Park: Operation Genesis
- Jurassic World: The Game

2018 video game developed by Frontier Developments

Video games developed in the United Kingdom


Crash: Mind over Mutant

Crash: Mind over Mutant is a platform video game developed by Radical Entertainment for the PlayStation 2, PlayStation Portable (ported by Virtuos), Wii and Xbox 360. The Nintendo DS version of the game was developed by TOSE. It was released in North America on October 7, 2008 and was later released in Europe and Australia on October 29 and October 31 respectively. It is the second game in the series not to have a Japanese release, after Crash of the Titans. It is the first game in the franchise to be published by Activision following its merging with the series' previous publisher Vivendi Games, and the last major console entry as a whole until 2017's Crash Bandicoot N. Sane Trilogy, a remaster of the first three Crash games.

Crash: Mind over Mutant is the fifteenth installment in the Crash Bandicoot video game series, and the seventh in the main franchise. The game's story centers on the arrival of a popular technological device (a parody of devices such as the iPhone and BlackBerry) that puts whoever uses it under the control of the device's creators, Doctor Neo Cortex and Doctor Nitrus Brio. Crash Bandicoot – protagonist of the series and only one unaffected by the device besides his spirit ally Aku Aku – must free his friends from the control of the device and put an end to Cortex and Brio's plot.

The title received an overall mixed response, with opinions varying from version to version. The PlayStation 2 and Wii versions were met with generally favourable reviews, while the Xbox 360 version generated a less positive reaction. Much praise was reserved for the voice acting and the satirical humor of the game's narrative, cutscenes and dialogue, while the unadjustable camera and extensive backtracking were heavily criticised. The Nintendo DS version was received negatively for what critics described as dull and repetitive gameplay.

## Gameplay

Mind over Mutant is a beat-'em-up game in which the player controls Crash Bandicoot, whose main objective is to explore his home, Wumpa Island, and surrounding environments to uncover the mystery behind the "NV", a personal digital assistant created by the main antagonist. Crash: Mind over Mutant is a mission-based game as opposed to Crash of the Titans's level-based structure, with goals being given by non-playable characters. Characters with exclamation points (!) over their heads, when talked to, give out important information that is needed to continue the story.

Scattered throughout the world are special items that empower Crash in one way or another. Golden stopwatches trigger a time-limited minigame which usually involves collecting objects or breaking things to earn bonuses. Red running shoes serve to temporarily power up Crash's kick attack and cause him to move faster, while the "Quad Damage" power-up can be used by Crash to temporarily multiply his attack power by four.

### Combat

Crash can perform both a light and heavy-powered attack. He can also block, dodge-and-counter, or break an enemy's block. When he defeats enemies or destroys objects, a magical substance known as Mojo is released that, when collected, allows Crash's abilities to be upgraded.

While small minions require only a single combo attack for Crash to defeat, larger enemies, known as "Titans", require more effort to subdue. Each of the unique Titans in the game possess a star meter that indicates how close they are to being stunned. When the meter is full, the Titan is susceptible to "jacking", meaning Crash can mount the creature and control it. While controlled by the player, the Titan possesses a similar moveset to Crash, although some jacked Titans can shoot projectiles. Besides having more health, the Titans controlled by Crash have a purple Titan Meter which, when full, allows the Titan to unleash a powerful special attack. This fully drains the meter in the process.

A feature new to the series involves Crash storing a Titan in his pocket and utilizing it when it is most advantageous. When Crash is riding a Titan, any Mojo collected goes to the Titan. If enough is collected, the Titan's abilities are upgraded rather than Crash's.

### Co-operative play

A player using a second controller can join in the game at any time. The second player will initially appear as a mask floating around Crash. The mask can attack enemies by using magic projectiles. By pressing a certain button, the second player's character can exit and enter the mask. In the Wii and Xbox 360 versions of the game, Crash's sister, Coco Bandicoot, fills in the role of the second player, while a white-furred version of Crash known as "Carbon Crash" takes on this role in the PlayStation 2 version. Coco is not available for play until she is defeated in a boss fight, after which she is fully playable (though the player can choose to continue with Carbon Crash). This method of play is useful for overpowering enemies and collecting Mojo faster.

## Plot

### Setting

The game is played in a free-roaming format rarely seen in previous games, although the DS version is played as a 2-D beat 'em up as a means of utilising the handheld's hardware to its greatest capacity. The opening events of the game take place on Wumpa Island. Crash's house is accessible here, and can be used by the player to access skins, concept art, enemy bios and cutscenes. Surrounding environments consist of societies inhabited by friendly Titans, including the Ratcicle Kingdom, the Rhinoroller Desert and the Sludge Junkyard. Other areas include the Evil Public School, attended by the main antagonist's niece, and Mount Grimly, where the evil mask Uka Uka is held. The final events of the game take place in the Space Head, a space station recycled from junkyard parts that is operated by Doctor Neo Cortex.

### Characters

Nine returning characters from previous Crash titles star in the home console version of Crash: Mind over Mutant. The protagonist of the game, Crash Bandicoot, is a bandicoot who must defeat the main antagonist Doctor Cortex and free the Titans from the control of Cortex's new device. Aiding Crash is Aku Aku, an ancient wooden mask who can take control of Titans by entering their heads. Crash's genius sister, Coco Bandicoot, appears initially as a boss character under the control of Cortex's and Brio's handheld device. Afterward, she acts as a playable character during the co-operative mode of the Wii and Xbox 360 console versions; she is replaced by a white-furred version of Crash named "Carbon Crash" in the PlayStation 2 version. Crunch Bandicoot, Crash's muscular cyborg friend, also appears in the game as a brainwashed boss.

The main antagonist of the series, Doctor Neo Cortex, is a mad scientist who plans on taking control of the denizens of the Wumpa Islands by marketing a mind-controlling mobile phone, the "NV", to them. Doctor Nitrus Brio makes a return appearance as Cortex's partner, aiding him in the development of the NV. Uka Uka, Aku Aku's evil twin brother, is the source of the bad Mojo necessary to take control of those using the NV device. Nina Cortex, Neo Cortex's niece, also appears in the game, having been transferred to an evil public school by her uncle. Doctor N. Gin, Cortex's demented right-hand man, makes a relatively minor appearance as a boss character early in the game.

A number of small fodder enemies attempt to hinder Crash in his journey, much of them appearing in the previous Crash of the Titans. The Ratnicians, who previously worked for Doctor Neo Cortex, have since gone feral and vegetarian, and now work for the minor antagonist Doctor N. Gin. The Brat Girls, who previously worked for Nina Cortex, have betrayed her and now run the Evil Public School and supervise the Ratcicles in the Ratcicle Kingdom. The Doom Monkeys make a return appearance as employees of Nitrus Brio in the Sludge Junkyard. Two new fodder enemies make their debut in the game: the Znu, a group of small, warty creatures that inhabit Mount Grimly, and the Slap-E's, hand-shaped robots that share Doctor Cortex's memories and personal problems.

### Story

Social satire and the theme of consumerism are frequent sources of humor in the plot of Crash: Mind over Mutant, with the game featuring jokes about SUVs and the skyrocketing prices of gas at the time. The game's story is told through a number of cut scenes animated in 2D Animation with different styles, such as those of Dragon Ball, The Animatrix, and South Park.

Crash: Mind over Mutant takes place a year later after the events of where Crash of the Titans left off. The Titans, free from the control of Doctor Neo Cortex, have spread throughout the Wumpa Islands and cultivated their own societies and towns. All seems well until a rejuvenated Cortex teams up with his old partner Nitrus Brio to invent a personal digital assistant, the "NV", that can control the minds of both mutants and bandicoots by transmitting bad Mojo, forcefully siphoned from Cortex's former boss Uka Uka. Crash and Aku Aku are not affected by the device, meanwhile N. Gin launches an attack on the bandicoots. Crash and Aku Aku defeated him, but they discover their family members Coco and Crunch are soon transformed into monstrous version of themselves who seek Crash's destruction.

After Coco is freed from the NV's control, the Bandicoots learn of Doctor Cortex's plot by accessing his blog, and decide to go to the Evil Public School, where Nina was placed by Doctor Cortex. Nina in return for Crash saving her science fair project, informs the heroes of Cortex's and Brio's alliance and reveals their location in the Junkyard, where N. Brio is recycling parts to create a new space station, the "Space Head", for Cortex and new NV devices.

Crash and Aku Aku find Crunch and N. Brio at the Junkyard and manage to break Crunch free from the NV's control. After learning of Uka Uka's whereabouts from N. Brio, Crash and Aku Aku go to Mount Grimly, where Uka Uka is being drained of all his Mojo. After being freed, Uka Uka informs Crash that his six voodoo bones, his source of power, have been stolen and given to three of the worst Titans on the Island, and tells them to go and retrieve the bones for him. When the bones have been gathered, Uka Uka uses his restored power to send Crash to the Space Head and Crash can enact Uka Uka's vengeance on Cortex. Crash engages in a fight against Cortex, who uses a mutation formula stolen from Brio to empower himself for the battle. Upon losing to the Bandicoot, Cortex throws a tantrum, causing the Space Head to fall towards the Earth. Cortex, having returned to normal, makes his escape from the plummeting Space Head, while Crash and Aku Aku brace themselves for a crash-landing on Wumpa Island. Crash and Aku Aku manage to survive the crash and reunite with Coco and Crunch as parts of the Space Head rain down from the sky. Coco ends the story by telling Aku Aku that they're not going to help clean up, much to his chagrin.

## Development and release

Development on Crash: Mind over Mutant began immediately after the completion of Crash of the Titans. The idea of preserving a titan for later use came from the play testing sessions of Crash of the Titans, in which the testers were found to be reluctant to leave the titans behind after an epic battle was won. Fans of the series were also a source of inspiration for Crash: Mind over Mutant, having such wishes as a free-roaming environment, Coco Bandicoot being a playable character and the return of the character Doctor Nitrus Brio. Full camera control was considered for the game, but was rejected for graphical reasons and to avoid having to insert a split-screen view in the cooperation mode. Online gameplay was also considered as a feature in the finished game, but was omitted due to the brief development schedule. Coco Bandicoot as a playable character was omitted from the PlayStation 2 version of the game due to her distinct animations taking up much of the console's memory. The Wii version of Crash: Mind over Mutant was created first, with the graphics scaled up for the Xbox 360, and scaled down for the PlayStation 2. A PlayStation 3 version of the game was rumored, but was promptly debunked by Radical Entertainment as a mistake on many press sites' behalf.

There was an open call for fan art of Crash Bandicoot to be submitted as part of a contest. The contest was hosted by Kidzworld as part of a preview page and was aimed at fans under the age of 18. Selected artwork is included in the final build of the game either inside of a comic book in Crash's house or on a wall in the school attended by Nina Cortex. In addition, winners had their names appear in the game's credits and received a free copy of the game when it was released.

To ensure that the audio did not become repetitive, Crash: Mind over Mutant features more than 8,500 lines of dialogue. The music was composed by Marc Baril, who previously composed Crash Tag Team Racing and Crash of the Titans.

## Reception

The console versions of Crash: Mind over Mutant received "mixed or average" reviews according to Metacritic. Dakota Grabowski of GameZone considered the game "slightly better" than Crash of the Titans, noting improvements in controls, gameplay and storyline from its predecessor, but lamented that "the camera almost ruins the whole package." Louis Bedigian of GameZone stated that the young demographic of the Crash Bandicoot series "may be annoyed by the camera and/or repetitive objectives but will be entertained by what is the best Crash game developed in a long time – and one of the best Mario clones released for Nintendo Wii." Neal Ronaghan of Nintendo World Report, meanwhile, felt that "the charming story and genuine humor" of the game were "overshadowed by the shallow gameplay". Christopher Ewen of GameZone praised the series' transition into a free-roaming gameplay environment, but felt that the game was too easy. Matt Casamassina of IGN found the game to be "exactly like its predecessor: a mildly enjoyable beat 'em up romp that's sure to satisfy younger players, but won't provide much that seasoned players haven't already experienced before". Andy Eddy of TeamXbox noted the uninspired gameplay, which was "not very consistent in its fun generation." Justin Calvert of GameSpot claimed that the game "has too much backtracking and too many camera problems to make it recommendable". Dan Pearson of Eurogamer criticised the game's fixed camera, extensive backtracking, useless mission log and dated concepts. Andrew Reiner of Game Informer proclaimed that the game's "unpredictable framerates, extensive backtracking, and the guarantee of one death caused by faulty camera positioning for every platforming segment" made it "a mess of a game, and a new low point for the once-loved marsupial."

Grabowski and Ewen dismissed the in-game graphics as run-of-the-mill, with Grabowski elaborating that while the animations have improved from Crash of the Titans, the "semi-decent" environments have not. Towell and Casamassina, however, were impressed by the large draw distances, detailed environments, and special effects such as transparent water and reflective ice. The frame rate was said to suffer from slowdown during extensive action and especially on weaker consoles, whereas Towell regarded the frame rate as solid. The cutscenes were singled out for praise for their varied art styles, though Eddy was disconcerted by the abrupt transitions between gameplay and cutscenes. Bedigian regarded the overall visuals as decent, but was critical of Crash's "dorky" design, feeling that the subtle changes that have been made over the years "really hurt Crash’s appearance as a leading game character".

Critics lauded the humorous voice-acting and dialogue, with Towell characterizing the game as a well-done mimicry of Saturday-morning cartoons. However, Casamassina found some of the in-game dialogue annoying and repetitive; the music was similarly described as such by Towell, Bedigian and Grabowski, while Ewen regarded it as "standard Crash fare".

Reviews for the Nintendo DS version were generally unfavourable, with most reviewers criticizing the level design for its lack of challenge and inspiration. Mike David of GameZone found it to be the first Crash title to disappoint him; he derided the confusing storyline and subpar audio, and while he found the character models to be clean and polished, he noted that the frame rate suffered from stuttering and slowdown. Mike Cook of Pocket Gamer appreciated the game's sense of "good, clean fun" and replayability granted by its optional collectibles, minigames and unlockable content, but felt that the power of the mutants made the gameplay "strangely effortless" and led to a disappointing campaign length. He also described the graphics as "splotchy" and considered them outdated and simplistic compared to the Final Fantasy IV remake. Neal Ronaghan of Nintendo World Report deemed the game to be a competent and enjoyable platformer in short bursts, but disliked the emphasis on piloting mutants as it diverted focus from the game's titular character. Craig Harris of IGN interpreted the developer's lack of credit in the game as embarrassment for having been involved in its production, and he condemned the game's "lifeless" execution in comparison to both Amaze Entertainment's version of Crash of the Titans on the same system and the Game Boy Advance installments by Vicarious Visions.

2008 video game