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The Divergent Series
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For the book series on which the film series is based, see Divergent trilogy. For the mathematical use, see Divergent series.
The Divergent Series
Produced by
Lucy Fisher
Douglas Wick
Pouya Shabazian
Based on
The Divergent trilogy
by Veronica Roth
Starring
Shailene Woodley
Theo James
Zoë Kravitz
Miles Teller
Ansel Elgort
Production
company
Red Wagon Entertainment
Summit Entertainment
Distributed by
Summit Entertainment
Lionsgate
Release date
1: March 21, 2014 (2014-03-21)
2: 000000002015-03-20-0000March 20, 2015
3: 000000002016-03-18-0000March 18, 2016
4: TBA
Running time
379 minutes (3 films)
Country
United States
Language
English
Budget
Total (3 films):
$305 million
Box office
Total (3 films):
$765,409,015
The Divergent Series is a feature film trilogy based on the Divergent novels by the American author Veronica Roth. Distributed by Summit Entertainment and Lionsgate Films, the series consists of three science fiction films set in a dystopian society. They have been produced by Lucy Fisher, Pouya Shabazian, and Douglas Wick.
The series star Shailene Woodley and Theo James as lead characters Beatrice Prior (Tris) and Tobias Eaton (Four), respectively. The supporting cast includes Ansel Elgort, Zoë Kravitz, and Miles Teller. Kate Winslet played the main antagonist in the first two films. The first film in the series was directed by Neil Burger, while the second and third films were directed by Robert Schwentke.
The Veronica Roth novels consist primarily of the trilogy ('Divergent', 'Insurgent', 'Allegiant'). Development began in 2011 following Summit's acquisition of the film rights to the Divergent novel in partnership with production company Red Wagon Entertainment.[1] The studios announced production on the sequel following the first film's strong performance in Thursday late-night screenings, where it grossed $4.9 million.[2] They acquired film rights to the Allegiant novel in December 2013, deciding in April 2014 to split the third novel into a two-part film adaptation.[3]
The first installment, Divergent (2014), grossed over $288 million worldwide,[4] while the second installment, The Divergent Series: Insurgent (2015), grossed over $297 million worldwide.[5] Insurgent was also the first Divergent film to be released in IMAX 3D.[6] The third installment, The Divergent Series: Allegiant (2016), grossed $179 million. Thus, the first three films of the series have grossed over $765 million worldwide. A fourth film, The Divergent Series: Ascendant was to be released theatrically, but due to Allegiant's poor showing at the box office, it was announced it will be released as a TV movie.
Contents
[hide]
1 Development
1.1 Casting the main roles
2 Production
2.1 Directors
2.2 Scripts
2.3 Cast
2.4 Filming
2.5 Music
3 Plot
3.1 Divergent (2014)
3.2 The Divergent Series: Insurgent (2015)
3.3 The Divergent Series: Allegiant (2016)
3.4 Cancelled film
3.5 Ascendant (TV series)
4 Recurring cast and characters
5 Crew
6 Reception
6.1 Box office performance
6.2 Critical and public response
7 See also
8 References
9 External links
Development[edit]
In March 2011, Summit Entertainment picked up the rights for Divergent with Douglas Wick and Lucy Fisher's production company Red Wagon Entertainment, while Lionsgate distributes the franchise.[1] Neil Burger was announced as the director on August 23, 2012.[7] Evan Daugherty, who co-wrote the screenplay with Vanessa Taylor, said, "I get hung up on the toughness of the movie but of equal importance is the love story between Tris and Four. It's inherently and inextricably linked to Tris' character journey. There will be plenty of sexual tension and chemistry, but it's important that all of that stuff doesn't just feel like it's thrown in, but that it all helps Tris grow as a character."[8] Daugherty further added, "It's tricky because the book is a very packed read with a lot of big ideas. So, distilling that into a cool, faithful two-hour movie is challenging. Not only do you have to establish five factions, but you have to acknowledge that there's a sixth entity, which is the divergent, and you also have the factionless. So there's a world that really has to be built out for the big screen… the movie is going to do it a little more efficiently."[9] Author Veronica Roth said about the script of the film: "Reading a script is a really interesting experience. I'd never read a script before. I was really impressed by how closely it stuck to the general plot line of the book."[10]
Initially, the budget of the film was $40 million but later Lionsgate increased it to $80 million (which finally changed to $85 million) due to the success of The Hunger Games. Analyst Ben Mogil said, "Divergent is more similar to Hunger Games in that the company owns the underlying economics (i.e. production) and the budget (at $80[million]) is more manageable."[11]
In December 2013, Summit Entertainment announced that a film adaptation of Insurgent, the second novel in the Divergent trilogy, would be released as The Divergent Series: Insurgent on March 20, 2015,[12] as a sequel to the film adaptation of Divergent with Brian Duffield originally chosen to write the script for the film.[13][14] On December 16, 2013, it was announced that Neil Burger, director of Divergent, would not return to direct Insurgent as he was still working on the first film.[15] On February 13, 2014, it was announced that Robert Schwentke was offered the director position for the film and that Akiva Goldsman had been hired to re-write Duffield's script.[16][17]
Also in December 2013, Summit Entertainment announced that the film adaptation of Allegiant, the third and final novel in the Divergent trilogy, would be released in March 2016, serving as the finale of the series, which at the time was planned as a trilogy,[12] but on April 11, 2014, the studio decided to split the novel into a two-part film, much the like the Harry Potter, Twilight, and Hunger Games franchises did with the finales of their series. Lionsgate Motion Picture Group co-chairmen Rob Friedman and Patrick Wachsberger said in a statement that, "Veronica Roth brings her captivating story to a masterful conclusion in 'Allegiant,' a rich, action-packed book with material that is ideally suited to two strong and fulfilling movies. The storytelling arc and world of the characters lend themselves perfectly to two films, a storytelling strategy that has worked very well for us on the two Twilight Breaking Dawn films and about which we're tremendously enthusiastic for the two upcoming Mockingjay films of The Hunger Games franchise".[18] Noah Oppenheim was announced as the screenwriter for the first part of the Allegiant adaptation on July 9, 2014.[19] On December 5, 2014, it was announced that Robert Schwentke will return to direct Part 1.[20] On September 10, 2015, it was announced that the two films would be re-titled, with Part 1 being renamed as The Divergent Series: Allegiant and Part 2 as The Divergent Series: Ascendant.[21]
Casting the main roles[edit]
Shailene Woodley (left) and Theo James (right) at the Divergent LA Premiere on March 18, 2014.
On October 22, 2012, it was announced that Shailene Woodley had landed the lead role of Beatrice "Tris" Prior.[22] Lucas Till, Jack Reynor, Jeremy Irvine, Alex Pettyfer, Brenton Thwaites, Alexander Ludwig and Luke Bracey were all considered for the role of Tobias "Four" Eaton.[23][24] On March 15, 2013, it was announced that Theo James had been cast as Four.[25]
Though James was 10 years older than the character when cast, Roth praised his casting "I was sure within seconds: this was 'Four', no question. Theo is able to capture 'Four's' authority and strength, as well as his depth and sensitivity." She also mentions the chemistry between him and Shailene: "He is a perfect match for Shailene's incredibly strong presence as Tris. I'm thrilled!"[26] The producers said about his casting: "We took our time to find the right actor to fill the role of Four, and Theo is definitely the perfect fit. Veronica has crafted a truly iconic character in Four and we cannot wait to begin production and bring him and this story to life for millions of fans around the world."[25]
Production[edit]
Filming for the franchise began on April 16, 2013.
Film
Producers
Director
Screenwriters
Novel by Veronica Roth
Divergent Fisher , Lucy Lucy Fisher
Douglas Wick
Pouya Shabazian Burger , Neil Neil Burger Evan Daugherty
Vanessa Taylor Divergent
The Divergent Series: Insurgent Schwentke , Robert Robert Schwentke Akiva Goldsman
Brian Duffield Insurgent
The Divergent Series: Allegiant Noah Oppenheim
Adam Cooper
Bill Collage Allegiant
The Divergent Series: Ascendant Lee Toland Krieger
Lucy Fisher, Douglas Wick, and Pouya Shabazian have been producers for all four films in the series. Veronica Roth and Neil Burger have acted as executive producers on the first two films. Summit Entertainment and Lionsgate collaborated on all four films. On April 11, 2014, the studio decided to split the final book, Allegiant, into two parts, much like Harry Potter and the Deathly Hallows – Parts 1 and 2, The Twilight Saga: Breaking Dawn – Parts 1 and 2, and The Hunger Games: Mockingjay – Parts 1 and 2.
Directors[edit]
Neil Burger directed the first film, Divergent, and was originally going to direct the sequel, The Divergent Series: Insurgent, but in December 2013 it was announced that Burger would not return, due to him still working on post-production work for the first film. Speaking on the subject the director stated, "There was no breathing room. They were like, 'We're holding to a March 2015 release date, and we have to start no later than May 2014.' It was a recipe for failure, at least with me on board. I wouldn't be able to do either thing right. I reluctantly let it go".[27] In February 2014, it was confirmed that Robert Schwentke would direct the sequel instead.[16] On December 5, 2014, it was announced that Schwentke would return for The Divergent Series: Allegiant.[28]
Scripts[edit]
Snow White and the Huntsman screenwriter Evan Daugherty and Hope Springs screenwriter Vanessa Taylor co-wrote the screenplay for Divergent. During the time that Neil Burger was originally going to direct the sequel, Brian Duffield was recruited to write the screenplay for The Divergent Series: Insurgent, but after Burger dropped out of the film and Robert Schwentke took over as director, he brought in Akiva Goldsman and Mark Bomback to rewrite Duffield's script.
The screenplay for the two-part finale, The Divergent Series: Allegiant and Ascendant, was written by Noah Oppenheim, Adam Cooper, Bill Collage, and Stephen Chbosky.
Cast[edit]
On March 11, 2013, it was announced that the roles of Tori, Christina, and Caleb had gone to Maggie Q, Zoë Kravitz, and Ansel Elgort, respectively.[29] Ray Stevenson, Jai Courtney and Aaron Eckhart were announced to be in talks to join the cast on March 15, 2013,[30] Stevenson and Courtney joined the cast as Marcus Eaton and Eric, respectively.[31] That same day, Miles Teller was cast as Peter.[32] The studio’s announcement included a bit of casting news: Mekhi Phifer will play Dauntless leader Max. Kate Winslet was announced to be in talks on January 24, 2013.[33] Later it was confirmed that she would portray Jeanine Matthews.[34] Talking about playing the negative character for the first time, Winslet said, "I'm no idiot. The idea went through my head that I have never played a baddie before, I was almost kind of surprised." Since Winslet joined the filming late, she used that distance from her co-stars to appear aloof on the first day of her shoot. "I wanted to break it and say, 'It's OK, I'm really fun. I promise.' But I thought, just for today, I'd let them think that I am a complete bitch."[35] On March 25, 2013, Ben Lamb was cast as Edward, Ben Lloyd-Hughes as Will, and Christian Madsen as Al.[36] In April 2013, Ashley Judd and Tony Goldwyn joined the cast as Natalie and Andrew Prior, Beatrice's parents.[37][38]
In 2014, the cast for the second film was announced. On May 12, 2014, it was announced that Octavia Spencer joined the cast as Amity representative Johanna Reyes.[39] Late May 2014, Suki Waterhouse and Jonny Weston were cast as Marlene and Edgar respectively.[40] Early June 2014, Stephanie Leigh Schlund announced that she was cast in the film as a member of the Amity,[41] and Naomi Watts and Daniel Dae Kim joined the cast as Evelyn Johnson and Jack Kang.[42][43] On June 9, 2014, Rosa Salazar joined the cast as Lynn.[44] On June 10, 2014, Australian actor Keiynan Lonsdale joined the cast as Uriah.[45] On June 11, 2014, Emjay Anthony joined the cast as Hector.[46]
On April 28, 2015, it was reported that Golden Globe nominee Jeff Daniels joined the cast for Allegiant as David, the leader of the Bureau of Genetic Welfare.[47] On May 1, 2015, it was announced that Bill Skarsgård had joined the cast as Matthew.[48]
Filming[edit]
Principal photography of Divergent began on April 16, 2013, and concluded on July 16, 2013. Almost all of the filming took place in Chicago with reshoots taking place from January 24–26, 2014, in Los Angeles.
Principal photography for The Divergent Series: Insurgent began on May 27, 2014, in Atlanta, Georgia[49] and concluded on September 6, 2014. Filming took place at United States Penitentiary, Atlanta. A set was constructed for the Amity Compound at Serenbe Community South of Atlanta. In downtown Atlanta, a zip-line scene was filmed for which a set was constructed on the roof of Peachtree Center. For two days, production was moved to Chicago with scenes being filmed in Wells Street, Franklin Street, and Adams Street. Production was then moved back to Atlanta for reshoots.
Principal photography for The Divergent Series: Allegiant began in Atlanta on May 18, 2015,[50][51] with filming concluding on August 23, 2015.
Music[edit]
Junkie XL scored the first film in the film series, Divergent, while Joseph Trapanese scored the second and third entries, Insurgent and Allegiant.
Plot[edit]
Divergent (2014)[edit]
Main article: Divergent (film)
In a world where people are divided into distinct factions based on human virtues, Tris Prior is warned she is Divergent and will never fit into any one group. When she discovers a conspiracy by a faction leader to destroy all Divergents, Tris must learn to trust in the mysterious Four and together they must find out what makes being Divergent so dangerous before it's too late.
The Divergent Series: Insurgent (2015)[edit]
Main article: The Divergent Series: Insurgent
As they search for allies and answers in the wake of the uprising, Tris and Four are on the run. Hunted by Jeanine Matthews, leader of the Erudite faction, Tris and Four race against time as they try to figure out what Abnegation sacrificed their lives to protect, and why the Erudite leaders will do anything to stop them. Haunted by her past choices but desperate to protect the ones she loves, Tris faces one impossible challenge after another as she unlocks the truth about the past and ultimately the future of her world.
The Divergent Series: Allegiant (2016)[edit]
Main article: The Divergent Series: Allegiant
On December 16, 2013, Summit Entertainment announced that the film adaptation of Allegiant would be released on March 18, 2016.[12] On April 11, 2014, Lionsgate announced that the film adaptation would be split into two films with the first part titled, The Divergent Series: Allegiant – Part 1.[52] The film was then re-titled as The Divergent Series: Allegiant.[21]
Tris must escape with Four and go beyond the wall enclosing Chicago. For the first time ever, they will leave the only city and family they have ever known in order to find a peaceful solution for their embroiled city. Once outside, old discoveries are quickly rendered meaningless with the revelation of shocking new truths. Tris must quickly decide who she can trust as a ruthless battle ignites beyond the walls of Chicago. In order to survive, Tris will be forced to make impossible choices about courage, allegiance, sacrifice and love.
Cancelled film[edit]
A Third sequel to the film was initially planned, with a March 2017 release, based on the latter half of the Allegiant book.[53] The timing was changed to a June 2017 release, with Lee Toland Krieger then attached as the movie's director.[54][55]
Ascendant (TV series)[edit]
In July 2016, after the poor performance of the third film at the box office, the producers considered creating the fourth as a television film, which would add new characters to the story who could continue in a spinoff television series, moving beyond the books.[54][56]
In September 2016, Shailene Woodley stated on Today that the film versus television decision was not finalized, and that it was "a limbo waiting game".[57] In the same interview Woodley spoke despairingly of the chances of her returning to the project in a television format, although noting she would be open to returning to it as a theatrical film.[57] In February 2017, with the producers having confirmed that it would be a television project, it was announced that Woodley had backed out of her starring role.[58]
In August 2017, Starz and Lionsgate Television announced that they were beginning to develop the TV series.[59]
Recurring cast and characters[edit]
See also: List of Divergent characters
Divergent
(2014)
The Divergent Series:
Insurgent
(2015)
The Divergent Series:
Allegiant
(2016)
The Divergent Series:
Ascendant
(TBA)
Beatrice "Tris" Prior
Shailene Woodley TBA
Tobias "Four" Eaton
Theo James TBA
Christina
Zoë Kravitz TBA
Peter Hayes
Miles Teller TBA
Caleb Prior
Ansel Elgort TBA
Marcus Eaton
Ray Stevenson
Tori Wu
Maggie Q
Jeanine Matthews
Kate Winslet
Natalie Prior
Ashley Judd
Eric Coulter
Jai Courtney
Max
Mekhi Phifer
Will
Ben Lloyd-Hughes
Andrew Prior
Tony Goldwyn
Lauren
Justine Wachsberger
Evelyn Johnson-Eaton
Naomi Watts TBA
Johanna Reyes
Octavia Spencer
Uriah Pedrad
Keiynan Lonsdale
Edgar
Jonny Weston
David
Jeff Daniels TBA
Matthew
Bill Skarsgård TBA
Al
Christian Madsen
Molly
Amy Newbold
Edward
Ben Lamb
Jack Kang
Daniel Dae Kim
Marlene
Suki Waterhouse
Lynn
Rosa Salazar
Hector
Emjay Anthony
Edith Prior
Janet McTeer
Crew[edit]
Divergent
(2014)
The Divergent Series:
Insurgent
(2015)
The Divergent Series:
Allegiant
(2016)
The Divergent Series:
Ascendant
(TBA)
Director
Neil Burger Robert Schwentke Lee Toland Krieger
Producer(s)
Douglas Wick
Lucy Fisher
Pouya Shabazian N/A
Writer(s)
Evan Daugherty
Vanessa Taylor Brian Duffield
Akiva Goldsman
Mark Bomback Noah Oppenheim
Adam Cooper
Bill Collage
Based on
Divergent by Veronica Roth Insurgent by Veronica Roth Allegiant by Veronica Roth
Composer
Junkie XL Joseph Trapanese N/A
Cinematographer
Alwin H. Küchler Florian Ballhaus
Editor(s)
Richard Francis-Bruce
Nancy Richardson Nancy Richardson
Stuart Levy Stuart Levy N/A
Production companies
Red Wagon Entertainment
Summit Entertainment Red Wagon Entertainment
Summit Entertainment
Mandeville Films Red Wagon Entertainment
Mandeville Films N/A
Distributor
Lionsgate Summit Entertainment N/A
Running time
139 minutes 119 minutes 121 minutes TBA
Release date
March 21, 2014 March 20, 2015 March 18, 2016 TBA
Reception[edit]
Further information on the reception of each film: Divergent (film) § Reception, The Divergent Series: Insurgent § Reception, and The Divergent Series: Allegiant § Reception
Though the first two films were commercially successful, the series as a whole has not been well received by critics. Each successive film release has been rated lower than the previous release on review aggregator websites. Negative reviews of the third novel Allegiant and the studios' decision to split it into two films have been blamed for the third film's lackluster performance at the box office.
Box office performance[edit]
Film
Release date
Budget
Box office gross
Box office ranking
Ref(s)
North America
Other territories
Worldwide
All time
North America
All time
worldwide
Divergent March 21, 2014 (2014-03-21) $85 million $150,947,895 $137,937,923 $288,885,818 #283 #375 [60]
The Divergent Series: Insurgent March 20, 2015 (2015-03-20) $110 million $130,179,072 $167,097,257 $297,276,329 #392 #372 [5]
The Divergent Series: Allegiant March 18, 2016 (2016-03-18) $110 million $66,184,051 $113,062,817 $179,246,868 #1,088 #TBA [61]
Total
$305 million
$347,311,018
$418,097,997
$765,409,015
[62]
Critical and public response[edit]
Film
Rotten Tomatoes
Metacritic
CinemaScore
Divergent 41% (203 reviews)[63] 48 (38 reviews)[64] A[65]
The Divergent Series: Insurgent 28% (184 reviews)[66] 42 (40 reviews)[67] A-[68]
The Divergent Series: Allegiant 12% (171 reviews)[69] 33 (33 reviews)[70] B[71]
See also[edit]
2010s portal
Film in the United States portal
Science fiction portal
References[edit]
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Jump up ^ "The Divergent Series: Allegiant (2016)". Rotten Tomatoes. Flixster. Retrieved July 1, 2016.
Jump up ^ "The Divergent Series: Allegiant – Reviews (2016)". Metacritic. CBS Interactive.
Jump up ^ Anthony D'Alessandro (March 18, 2016). "'Allegiant' Posts Lowest Opening In 'Divergent Series': Was It Wise To Split The Last Novel Into Two Movies?". Deadline Hollywood. Retrieved March 19, 2016.
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Microevolution
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Microevolution is the change in allele frequencies that occurs over time within a population.[1] This change is due to four different processes: mutation, selection (natural and artificial), gene flow, and genetic drift. This change happens over a relatively short (in evolutionary terms) amount of time compared to the changes termed 'macroevolution' which is where greater differences in the population occur.
Population genetics is the branch of biology that provides the mathematical structure for the study of the process of microevolution. Ecological genetics concerns itself with observing microevolution in the wild. Typically, observable instances of evolution are examples of microevolution; for example, bacterial strains that have antibiotic resistance.
Microevolution over time leads to speciation or the appearance of novel structure, sometimes classified as macroevolution.[2] Macro and microevolution describe fundamentally identical processes on different scales.[2][3][4]
Contents
[hide]
1 Difference from macroevolution
2 Four processes
2.1 Mutation
2.2 Selection
2.3 Genetic drift
2.4 Gene flow
3 Origin and extended use of the term
3.1 Origin
3.2 Use in creationism
4 See also
5 References
6 External links
Difference from macroevolution[edit]
Macroevolution and microevolution describe fundamentally identical processes on different time scales.[5][2][page needed] Microevolution refers to small evolutionary changes (typically described as changes in allele frequencies) within a species or population.[6] while macroevolution is evolution on a scale of separated gene pools.[5] Macroevolutionary studies focus on change that occurs at or above the level of species.[7]
Four processes[edit]
Mutation[edit]
Main article: Mutation
Duplication of part of a chromosome
Mutations are changes in the DNA sequence of a cell's genome and are caused by radiation, viruses, transposons and mutagenic chemicals, as well as errors that occur during meiosis or DNA replication.[8][9][10] Errors are introduced particularly often in the process of DNA replication, in the polymerization of the second strand. These errors can also be induced by the organism itself, by cellular processes such as hypermutation.
During the process of DNA replication, errors occasionally occur in the polymerization of the second strand. These errors, called mutations, can affect the phenotype of an organism, especially if they occur within the protein coding sequence of a gene. Error rates are usually very low—1 error in every 10–100 million bases—due to the "proofreading" ability of DNA polymerases.[11][12] (Without proofreading error rates are a thousandfold higher; because many viruses rely on DNA and RNA polymerases that lack proofreading ability, they experience higher mutation rates.) Processes that increase the rate of changes in DNA are called mutagenic: mutagenic chemicals promote errors in DNA replication, often by interfering with the structure of base-pairing, while UV radiation induces mutations by causing damage to the DNA structure.[13] Chemical damage to DNA occurs naturally as well, and cells use DNA repair mechanisms to repair mismatches and breaks in DNA—nevertheless, the repair sometimes fails to return the DNA to its original sequence.
In organisms that use chromosomal crossover to exchange DNA and recombine genes, errors in alignment during meiosis can also cause mutations.[14] Errors in crossover are especially likely when similar sequences cause partner chromosomes to adopt a mistaken alignment making some regions in genomes more prone to mutating in this way. These errors create large structural changes in DNA sequence—duplications, inversions or deletions of entire regions, or the accidental exchanging of whole parts between different chromosomes (called translocation).
Mutation can result in several different types of change in DNA sequences; these can either have no effect, alter the product of a gene, or prevent the gene from functioning. Studies in the fly Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene, this will probably be harmful, with about 70 percent of these mutations having damaging effects, and the remainder being either neutral or weakly beneficial.[15] Due to the damaging effects that mutations can have on cells, organisms have evolved mechanisms such as DNA repair to remove mutations.[8] Therefore, the optimal mutation rate for a species is a trade-off between costs of a high mutation rate, such as deleterious mutations, and the metabolic costs of maintaining systems to reduce the mutation rate, such as DNA repair enzymes.[16] Viruses that use RNA as their genetic material have rapid mutation rates,[17] which can be an advantage since these viruses will evolve constantly and rapidly, and thus evade the defensive responses of e.g. the human immune system.[18]
Mutations can involve large sections of DNA becoming duplicated, usually through genetic recombination.[19] These duplications are a major source of raw material for evolving new genes, with tens to hundreds of genes duplicated in animal genomes every million years.[20] Most genes belong to larger families of genes of shared ancestry.[21] Novel genes are produced by several methods, commonly through the duplication and mutation of an ancestral gene, or by recombining parts of different genes to form new combinations with new functions.[22][23]
Here, domains act as modules, each with a particular and independent function, that can be mixed together to produce genes encoding new proteins with novel properties.[24] For example, the human eye uses four genes to make structures that sense light: three for color vision and one for night vision; all four arose from a single ancestral gene.[25] Another advantage of duplicating a gene (or even an entire genome) is that this increases redundancy; this allows one gene in the pair to acquire a new function while the other copy performs the original function.[26][27] Other types of mutation occasionally create new genes from previously noncoding DNA.[28][29]
Selection[edit]
Main articles: Natural selection and Artificial selection
Selection is the process by which heritable traits that make it more likely for an organism to survive and successfully reproduce become more common in a population over successive generations.
It is sometimes valuable to distinguish between naturally occurring selection, natural selection, and selection that is a manifestation of choices made by humans, artificial selection. This distinction is rather diffuse. Natural selection is nevertheless the dominant part of selection.
Natural selection of a population for dark coloration.
The natural genetic variation within a population of organisms means that some individuals will survive more successfully than others in their current environment. Factors which affect reproductive success are also important, an issue which Charles Darwin developed in his ideas on sexual selection.
Natural selection acts on the phenotype, or the observable characteristics of an organism, but the genetic (heritable) basis of any phenotype which gives a reproductive advantage will become more common in a population (see allele frequency). Over time, this process can result in adaptations that specialize organisms for particular ecological niches and may eventually result in the speciation (the emergence of new species).
Natural selection is one of the cornerstones of modern biology. The term was introduced by Darwin in his groundbreaking 1859 book On the Origin of Species,[30] in which natural selection was described by analogy to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favored for reproduction. The concept of natural selection was originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, nothing was known of modern genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical and molecular genetics is termed the modern evolutionary synthesis. Natural selection remains the primary explanation for adaptive evolution.
Genetic drift[edit]
Main article: Genetic drift
Ten simulations of random genetic drift of a single given allele with an initial frequency distribution 0.5 measured over the course of 50 generations, repeated in three reproductively synchronous populations of different sizes. In general, alleles drift to loss or fixation (frequency of 0.0 or 1.0) significantly faster in smaller populations.
Genetic drift is the change in the relative frequency in which a gene variant (allele) occurs in a population due to random sampling. That is, the alleles in the offspring in the population are a random sample of those in the parents. And chance has a role in determining whether a given individual survives and reproduces. A population's allele frequency is the fraction or percentage of its gene copies compared to the total number of gene alleles that share a particular form.[31]
Genetic drift is an evolutionary process which leads to changes in allele frequencies over time. It may cause gene variants to disappear completely, and thereby reduce genetic variability. In contrast to natural selection, which makes gene variants more common or less common depending on their reproductive success,[32] the changes due to genetic drift are not driven by environmental or adaptive pressures, and may be beneficial, neutral, or detrimental to reproductive success.
The effect of genetic drift is larger in small populations, and smaller in large populations. Vigorous debates wage among scientists over the relative importance of genetic drift compared with natural selection. Ronald Fisher held the view that genetic drift plays at the most a minor role in evolution, and this remained the dominant view for several decades. In 1968 Motoo Kimura rekindled the debate with his neutral theory of molecular evolution which claims that most of the changes in the genetic material are caused by genetic drift.[33] The predictions of neutral theory, based on genetic drift, do not fit recent data on whole genomes well: these data suggest that the frequencies of neutral alleles change primarily due to selection at linked sites, rather than due to genetic drift by means of sampling error.[34]
Gene flow[edit]
Main article: Gene flow
Gene flow is the exchange of genes between populations, which are usually of the same species.[35] Examples of gene flow within a species include the migration and then breeding of organisms, or the exchange of pollen. Gene transfer between species includes the formation of hybrid organisms and horizontal gene transfer.
Migration into or out of a population can change allele frequencies, as well as introducing genetic variation into a population. Immigration may add new genetic material to the established gene pool of a population. Conversely, emigration may remove genetic material. As barriers to reproduction between two diverging populations are required for the populations to become new species, gene flow may slow this process by spreading genetic differences between the populations. Gene flow is hindered by mountain ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which has hindered the flow of plant genes.[36]
Depending on how far two species have diverged since their most recent common ancestor, it may still be possible for them to produce offspring, as with horses and donkeys mating to produce mules.[37] Such hybrids are generally infertile, due to the two different sets of chromosomes being unable to pair up during meiosis. In this case, closely related species may regularly interbreed, but hybrids will be selected against and the species will remain distinct. However, viable hybrids are occasionally formed and these new species can either have properties intermediate between their parent species, or possess a totally new phenotype.[38] The importance of hybridization in creating new species of animals is unclear, although cases have been seen in many types of animals,[39] with the gray tree frog being a particularly well-studied example.[40]
Hybridization is, however, an important means of speciation in plants, since polyploidy (having more than two copies of each chromosome) is tolerated in plants more readily than in animals.[41][42] Polyploidy is important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to pair with an identical partner during meiosis.[43] Polyploid hybrids also have more genetic diversity, which allows them to avoid inbreeding depression in small populations.[44]
Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among bacteria.[45] In medicine, this contributes to the spread of antibiotic resistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other species.[46] Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzuki bean beetle Callosobruchus chinensis may also have occurred.[47][48] An example of larger-scale transfers are the eukaryotic bdelloid rotifers, which appear to have received a range of genes from bacteria, fungi, and plants.[49] Viruses can also carry DNA between organisms, allowing transfer of genes even across biological domains.[50] Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells and prokaryotes, during the acquisition of chloroplasts and mitochondria.[51]
Gene flow is the transfer of alleles from one population to another.
Migration into or out of a population may be responsible for a marked change in allele frequencies. Immigration may also result in the addition of new genetic variants to the established gene pool of a particular species or population.
There are a number of factors that affect the rate of gene flow between different populations. One of the most significant factors is mobility, as greater mobility of an individual tends to give it greater migratory potential. Animals tend to be more mobile than plants, although pollen and seeds may be carried great distances by animals or wind.
Maintained gene flow between two populations can also lead to a combination of the two gene pools, reducing the genetic variation between the two groups. It is for this reason that gene flow strongly acts against speciation, by recombining the gene pools of the groups, and thus, repairing the developing differences in genetic variation that would have led to full speciation and creation of daughter species.
For example, if a species of grass grows on both sides of a highway, pollen is likely to be transported from one side to the other and vice versa. If this pollen is able to fertilise the plant where it ends up and produce viable offspring, then the alleles in the pollen have effectively been able to move from the population on one side of the highway to the other.
Origin and extended use of the term[edit]
Origin[edit]
The term microevolution was first used by botanist Robert Greenleaf Leavitt in the journal Botanical Gazette in 1909, addressing what he called the "mystery" of how formlessness gives rise to form.[52]
..The production of form from formlessness in the egg-derived individual, the multiplication of parts and the orderly creation of diversity among them, in an actual evolution, of which anyone may ascertain the facts, but of which no one has dissipated the mystery in any significant measure. This microevolution forms an integral part of the grand evolution problem and lies at the base of it, so that we shall have to understand the minor process before we can thoroughly comprehend the more general one...
However, Leavitt was using the term to describe what we would now call developmental biology; it was not until Russian Entomologist Yuri Filipchenko used the terms "macroevolution" and "microevolution" in 1927 in his German language work, "Variabilität und Variation", that it attained its modern usage. The term was later brought into the English-speaking world by Theodosius Dobzhansky in his book Genetics and the Origin of Species (1937).[1]
Use in creationism[edit]
See also: Speciation
In young Earth creationism and baraminology a central tenet is that evolution can explain diversity in a limited number of created kinds which can interbreed (which they call "microevolution") while the formation of new "kinds" (which they call "macroevolution") is impossible.[3][53] This acceptance of "microevolution" only within a "kind" is also typical of old Earth creationism.[54]
Scientific organizations such as the American Association for the Advancement of Science describe microevolution as small scale change within species, and macroevolution as the formation of new species, but otherwise not being different from microevolution. In macroevolution, an accumulation of microevolutionary changes leads to speciation.[55] The main difference between the two processes is that one occurs within a few generations, whilst the other takes place over thousands of years (i.e. a quantitative difference).[56] Essentially they describe the same process; although evolution beyond the species level results in beginning and ending generations which could not interbreed, the intermediate generations could.
Opponents to creationism argue that changes in the number of chromosomes can be accounted for by intermediate stages in which a single chromosome divides in generational stages, or multiple chromosomes fuse, and cite the chromosome difference between humans and the other great apes as an example.[57] Creationists insist that since the actual divergence between the other great apes and humans was not observed, the evidence is circumstantial.
Describing the fundamental similarity between macro and microevolution in his authoritative textbook "Evolutionary Biology," biologist Douglas Futuyma writes,
One of the most important tenets of the theory forged during the Evolutionary Synthesis of the 1930s and 1940s was that "macroevolutionary" differences among organisms - those that distinguish higher taxa - arise from the accumulation of the same kinds of genetic differences that are found within species. Opponents of this point of view believed that "macroevolution" is qualitatively different from "microevolution" within species, and is based on a totally different kind of genetic and developmental patterning... Genetic studies of species differences have decisively disproved [this] claim. Differences between species in morphology, behavior, and the processes that underlie reproductive isolation all have the same genetic properties as variation within species: they occupy consistent chromosomal positions, they may be polygenic or based on few genes, they may display additive, dominant, or epistatic effects, and they can in some instances be traced to specifiable differences in proteins or DNA nucleotide sequences. The degree of reproductive isolation between populations, whether prezygotic or postzygotic, varies from little or none to complete. Thus, reproductive isolation, like the divergence of any other character, evolves in most cases by the gradual substitution of alleles in populations.
— Douglas Futuyma, "Evolutionary Biology" (1998), pp.477-8[2]
Contrary to the claims of some antievolution proponents, evolution of life forms beyond the species level (i.e. speciation) has indeed been observed and documented by scientists on numerous occasions.[58] In creation science, creationists accepted speciation as occurring within a "created kind" or "baramin", but objected to what they called "third level-macroevolution" of a new genus or higher rank in taxonomy. There is ambiguity in the ideas as to where to draw a line on "species", "created kinds", and what events and lineages fall within the rubric of microevolution or macroevolution.[59]
See also[edit]
Punctuated equilibrium - due to gene flow, major evolutionary changes may be rare
References[edit]
^ Jump up to: a b Microevolution: What is microevolution?
^ Jump up to: a b c d Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates.
^ Jump up to: a b edited by Scott, Eugenie C.; Branch, Glenn (2006). Not in our classrooms : why intelligent design is wrong for our schools (1st ed.). Boston: Beacon Press. p. 47. ISBN 0807032786. CS1 maint: Extra text: authors list (link)
Jump up ^ SA 80, http://www.scientificamerican.com/article/15-answers-to-creationist/
^ Jump up to: a b Matzke, Nicholas J. and Paul R. Gross. 2006. Analyzing Critical Analysis: The Fallback Antievolutionist Strategy. In Eugenie Scott and Glenn Branch, Not in Our Classrooms: Why Intelligent Design is Wrong for Our Schools, Beacon Press, Boston ISBN 0807032786[page needed]
Jump up ^ Reznick DN, Ricklefs RE (February 2009). "Darwin's bridge between microevolution and macroevolution". Nature. 457 (7231): 837–42. Bibcode:2009Natur.457..837R. PMID 19212402. doi:10.1038/nature07894.
Jump up ^ Dobzhansky, Theodosius Grigorievich (1937). Genetics and the origin of species. New York: Columbia Univ. Press. p. 12. LCCN 37033383.
^ Jump up to: a b Bertram J (2000). "The molecular biology of cancer". Mol. Aspects Med. 21 (6): 167–223. PMID 11173079. doi:10.1016/S0098-2997(00)00007-8.
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Jump up ^ Griffiths, Anthony J. F.; Miller, Jeffrey H.; Suzuki, David T.; Lewontin, Richard C.; Gelbart, William M., eds. (2000). "Spontaneous mutations". An Introduction to Genetic Analysis (7th ed.). New York: W. H. Freeman. ISBN 0-7167-3520-2.
Jump up ^ Freisinger, E; Grollman, AP; Miller, H; Kisker, C (2004). "Lesion (in)tolerance reveals insights into DNA replication fidelity.". The EMBO Journal. 23 (7): 1494–505. PMC 391067  . PMID 15057282. doi:10.1038/sj.emboj.7600158.
Jump up ^ Griffiths, Anthony J. F.; Miller, Jeffrey H.; Suzuki, David T.; Lewontin, Richard C.; Gelbart, William M., eds. (2000). "Induced mutations". An Introduction to Genetic Analysis (7th ed.). New York: W. H. Freeman. ISBN 0-7167-3520-2.
Jump up ^ Griffiths, Anthony J. F.; Miller, Jeffrey H.; Suzuki, David T.; Lewontin, Richard C.; Gelbart, William M., eds. (2000). "Chromosome Mutation I: Changes in Chromosome Structure: Introduction". An Introduction to Genetic Analysis (7th ed.). New York: W. H. Freeman. ISBN 0-7167-3520-2.
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Jump up ^ Sniegowski P, Gerrish P, Johnson T, Shaver A; Gerrish; Johnson; Shaver (2000). "The evolution of mutation rates: separating causes from consequences". BioEssays. 22 (12): 1057–66. PMID 11084621. doi:10.1002/1521-1878(200012)22:12<1057::AID-BIES3>3.0.CO;2-W. CS1 maint: Multiple names: authors list (link)
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