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Out of the Furnace
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Not to be confused with Out of This Furnace.
Out of the Furnace
Theatrical release poster
Directed by
Scott Cooper
Produced by
Jennifer Davisson Killoran
Leonardo DiCaprio
Ryan Kavanaugh
Ridley Scott
Michael Costigan
Written by
Brad Ingelsby
Scott Cooper
Starring
Christian Bale
Woody Harrelson
Casey Affleck
Forest Whitaker
Willem Dafoe
Zoë Saldana
Sam Shepard
Music by
Dickon Hinchliffe
Cinematography
Masanobu Takayanagi
Edited by
David Rosenbloom
Production
company
Appian Way Productions
Scott Free Productions
Red Granite Pictures
Distributed by
Relativity Media
Release date
December 6, 2013 (2013-12-06)
Running time
116 minutes
Country
United States
Language
English
Budget
$22 million[1]
Box office
$15.7 million[2]
Out of the Furnace is a 2013 American thriller film, directed by Scott Cooper, from a screenplay written by Cooper and Brad Ingelsby. Produced by Ridley Scott and Leonardo DiCaprio, the film stars Christian Bale, Casey Affleck, Woody Harrelson, Zoe Saldana, Forest Whitaker, Willem Dafoe, and Sam Shepard.
The film is about a Pennsylvania steel mill worker Russell Baze (Bale) and his Iraq war veteran brother Rodney (Affleck), who cannot adjust to civilian life. While Rodney makes some money doing bareknuckle fights for bar owner and small-time criminal John Petty (Dafoe), who runs illegal gambling operations, Rodney becomes so indebted due to his own gambling losses that he begs Petty to let him do a big money fight. After Petty reluctantly arranges for Rodney to do a fight for a ruthless criminal gang in the backwoods, Rodney disappears, and his brother tries to find out what has happened to him. The film received a limited release in Los Angeles and New York City on December 4, 2013, followed by a wide theatrical release on December 6. The film earned $15.7 million against its $22 million budget and it received mixed reviews.
Contents
[hide]
1 Plot
2 Cast
3 Production
3.1 Development
3.2 Filming
3.3 Music
4 Release
5 Reception
5.1 Box office
5.2 Critical response
5.3 Top ten lists
5.4 Lawsuit
6 References
7 External links
Plot[edit]
After getting off work at a North Braddock, Pennsylvania, steel mill, Russell Baze catches his brother Rodney at a horse racing simulcast, where Rodney had just bet on a losing horse. Rodney reveals John Petty loaned the money to him. Petty owns a bar and runs several illegal games. Russell visits Petty, pays off some of Rodney's debt, and promises to pay Petty the rest with his next paycheck if Rodney has not yet paid it off. Driving home intoxicated, Russell hits another car, killing its occupants, including a little boy. He is incarcerated for vehicular manslaughter. While in prison he is informed that his ailing father has died and his girlfriend Lena has left him for the small town police chief, Wesley Barnes. Upon his release from prison, Russell returns home and resumes his job at the mill.
The same day, Rodney participates in an illegal bare-knuckle prizefight. Rodney was supposed to take a "dive" for Petty as a way to repay some of the gambling debt he owes Petty, but during the fight, Rodney becomes enraged against his opponent and wins the fight. The next morning, Russell finds Rodney's bloodied knuckle tapes in the trash and confronts him about it. Russell wants him to work in the mill, but Rodney, a four-tour Iraq war veteran, is too mentally scarred to take a regular job.
Rodney tells Petty that these "nickel and dime" fights will never earn him enough money to pay Petty back. Rodney then insists that Petty call and organize a more lucrative fight. Petty reluctantly arranges a fight with Harlan DeGroat, a sociopathic drug dealer from rural New Jersey to whom Petty owes a lot of money. Russell wants Lena back, but she is pregnant with Wesley's baby. Russell unsuccessfully feigns happiness to Lena, saying she will be a great mom, but both know that her pregnancy makes their getting back together an impossibility.
Rodney is told he must purposely lose the fight in New Jersey. When DeGroat seeks assurances Rodney will lose, Petty promises he will. Rodney briefly knocks out his opponent, but when hearing Petty pleading with him, Rodney helps the fighter get up, takes a dive, and lets the man pummel his face into a bloody mess. After the fight, DeGroat asks for the rest of his money, but Petty reminds DeGroat they had agreed in advance that this fight made them even, and DeGroat drops the subject. While driving back home, DeGroat and his men ambush Petty and Rodney. DeGroat first shoots and kills Petty, has Rodney dragged into the woods, and kills him, too. Unknown to anyone, Petty had accidentally dialed his cell phone, which fell onto the car seat and connected to his bartender Dan's voicemail while recording DeGroat as clear evidence of his murdering Petty.
That night, Russell finds a letter from Rodney, stating this will be his last fight and he wants to work with Russell at the mill. Wesley informs Russell about Rodney's disappearance, and Russell and his uncle "Red" set off to find him. In DeGroat's town, Russell and Red are stopped by the sheriff, who informs them that DeGroat's men would kill them if they knew why the two were in town, and, as a favor to Sheriff Wesley, he will escort them to the state line rather than searching and arresting them for illegally carrying concealed weapons.
Upon returning to the mill, Wesley visits Russell and confirms Rodney's death. Russell goes to Petty's office, finds a phone number for DeGroat, and calls him without identifying himself, enticing him to come collect Petty's debt. At the bar, Russell sabotages DeGroat's van to prevent his escape and confronts him. DeGroat escapes to a nearby, shutdown mill, where Russell shoots him in the thigh. Russell then follows DeGroat to a field outside the mill as he hobbles off and shoots him in the back. Russell informs DeGroat that he is Rodney's brother, as Wesley approaches the field in a squad car. Wesley pleads for Russell to put down his gun, but Russell proceeds to carefully aim his hunting rifle and shoots DeGroat in the head. The film cuts to Russell sitting at home at the dining table and fades to black.
Cast[edit]
Christian Bale as Russell Baze
Woody Harrelson as Harlan DeGroat
Casey Affleck as Rodney Baze, Jr.
Forest Whitaker as Wesley Barnes
Willem Dafoe as John Petty
Tom Bower as Dan Dugan
Zoë Saldana as Lena Taylor
Sam Shepard as Gerald "Red" Baze
Production[edit]
Development[edit]
The film was produced by Relativity Media, Appian Way, Red Granite Pictures, and Scott Free Productions, with Jeff Waxman, Tucker Tooley, Brooklyn Weaver, Riza Aziz, Joey McFarland, Joe Gatta, Danny Dimbort, and Christian Mercuri serving as executive producers.[3] Director Scott Cooper read an article about Braddock, Pennsylvania, a declining steel industry town outside of Pittsburgh, and the efforts to revitalize it, led by mayor John Fetterman. After visiting, Cooper was inspired to use the borough as the backdrop for a film.[4] Cooper developed the story from The Low Dweller, a spec script written by Brad Ingelsby that had actor Leonardo DiCaprio and director Ridley Scott attached.[5] The studio offered the script to Cooper, which he rewrote, drawing on his experience of growing up in Appalachia and losing a sibling at a young age.[6] DiCaprio and Scott stayed on as producers of the film.[5] The story has no relation to Out of This Furnace, a 1941 historical novel by Thomas Bell, set in Braddock.[7] The Hollywood Reporter reported the film's budget was $22 million.[1]
Filming[edit]
Principal photography began in the Pittsburgh metropolitan area on April 13, 2012, and wrapped on June 1, 2012. The majority of filming took place in Braddock, and additional filming was in nearby North Braddock, Imperial, Rankin, and Swissvale.[8] Cinematographer Masanobu Takayanagi shot the feature in anamorphic format on Kodak 35 mm film.[9] Prison scenes were filmed in the Northern Panhandle of West Virginia, at the former State Penitentiary in Moundsville.[3] Filming also took place in rural Beaver County, including a deer hunting scene in Raccoon Creek State Park and a mill scene in Koppel.[10] Independence Township doubled for Bergen County, New Jersey.[11] The Carrie Furnace, an abandoned blast furnace near Braddock, served as the location for the film's finale.[4] Christian Bale wore a tattoo of Braddock's ZIP code, 15104, on his neck as a homage to the town's mayor John Fetterman, who has the same design on his arm.[12]
Music[edit]
The musical score to Out of the Furnace was composed by Dickon Hinchliffe. Originally, it was announced that Alberto Iglesias had reached an agreement to compose the score for the film.[13] However, Hinchliffe later took over scoring duties.[14] Pearl Jam frontman Eddie Vedder re-recorded the song Release from Pearl Jam's debut album Ten for the film, which can be heard during the opening scenes and end credits. A soundtrack album featuring Hinchliffe's score was released digitally on December 3, 2013 by Relativity Music Group.[15]
Release[edit]
The film premiered at the TCL Chinese Theatre on November 9, 2013 in Hollywood, California as part of the American Film Institute's AFI Fest.[16] It received a limited release in Los Angeles and New York City on December 4, 2013, followed by a wide theatrical release in the U.S. on December 6.[17] Director Scott Cooper won the award for best first or second film for Out of the Furnace at the 2013 Rome Film Festival.[18]
The Pearl Jam song "Release" is featured during the opening title and features in a newly recorded edition during the end credits.
Reception[edit]
Box office[edit]
Out of the Furnace was the only new film to receive a wide release in the U.S. on December 6, 2013, and earned an estimated $1.8 million on its opening day.[19] The film took in an estimated $5.3 million over its opening weekend.[20] The Los Angeles Times and The Hollywood Reporter characterized it as a box office bomb.[1][21] The film came in third behind the animated Disney film Frozen, which brought in $31.6 million, and The Hunger Games: Catching Fire, which had $27 million in ticket sales that weekend. Relativity Media had pre-sold the film to foreign distributors for $16 million, which offset its costs.[21]
Critical response[edit]
On Rotten Tomatoes, the film has an average approval rating of 53% based on 182 reviews and an average rating of 5.9/10. The general consensus for the site says: "While it may not make the most of its incredible cast, Out of the Furnace is still so packed with talent that it's hard to turn away."[22] The film holds a score of 63 (indicating "generally favorable reviews") out of 100 on Metacritic, based on 40 critics.[23]
Top ten lists[edit]
Despite its mixed critical reception, Out of the Furnace appeared on several critics' top ten lists of the best films of 2013.[24]
2nd – Richard Roeper, Chicago Sun-Times
2nd – Clint O'Connor, Cleveland Plain Dealer
4th – Marc Doye, Metacritic
6th – Chris Nashawaty, Entertainment Weekly
7th – Kristopher Tapley, Hitfix
Lawsuit[edit]
Town officials from Mahwah, New Jersey, urged a boycott of the film due to negative depictions of the Ramapough Mountain Indians, an indigenous people living around the Ramapo Mountains,[25] characterizing the film as a hate crime.[26] Relativity Media responded that the film "is not based on any one person or group" and is "entirely fictional".[27] Nine members of the group, eight of whom have the surname of the film's lead character, DeGroat, filed suit against the makers and other involved parties, claiming that Out of the Furnace portrays a gang of "inbreds" living in the Ramapo Mountains who are "lawless, drug-addicted, impoverished and violent."[28] The lawsuit asserts that "The Defendants, and each of them, knew or should have known that their actions would place Plaintiffs, and/or any person so situated in a false light." The suit continues, "The connection between the ethnic slur of 'Jackson Whites', with the location of the Ramapo Mountains of New Jersey', with a Bergen County Police patrol car, with the surnames 'DeGroat' and 'Van Dunk', is too specific to the Ramapough plaintiffs to be chance, coincidence or happenstance, and implies an element of knowledge on the part of the Defendants, or some of them."[29]
On May 16, 2014, U.S. District Court Judge William Walls, sitting in Newark, New Jersey, dismissed the lawsuit, saying that the film did not refer directly to any of the plaintiffs.[30]
References[edit]
^ Jump up to: a b c McClintock, Pamela (December 8, 2013). "Box Office: 'Frozen' Tops 'Catching Fire'; 'Out of the Furnace' DOA". The Hollywood Reporter. Retrieved December 8, 2013.
Jump up ^ "Out of the Furnace". Box Office Mojo. December 29, 2013. Retrieved June 17, 2016.
^ Jump up to: a b Wadas, Amy (April 13, 2012). "Moundsville Movie Great for Local Economy". WTRF. Retrieved April 16, 2012.
^ Jump up to: a b Corsaro, Louis A. (November 30, 2012). ""Out of the Furnace" shoot warm experience for Braddock". Pittsburgh Business Times. Retrieved July 12, 2013.
^ Jump up to: a b Fleming Jr., Mike (April 11, 2011). "‘Crazy Heart’s Scott Cooper Takes On ‘The Low Dweller’ For Relativity Media". Deadline.com. Retrieved November 30, 2013.
Jump up ^ Fleming Jr., Mike (November 20, 2013). "Fleming Q&As ‘Out Of The Furnace’ Helmer Scott Cooper". Deadline.com. Retrieved November 30, 2013.
Jump up ^ Corsaro, Louis A. (June 21, 2013). "Plot details emerge on 'Out of the Furnace'". Pittsburgh Business Times. Retrieved July 6, 2013.
Jump up ^ "Film Crews Prepare Braddock For 'Out Of The Furnace'". CBS Pittsburgh. April 13, 2012. Retrieved April 16, 2012.
Jump up ^ "ONFILM Interview: Masanobu Takayanagi". Kodak. October 15, 2013. Retrieved December 10, 2013.
Jump up ^ Tady, Scott (April 18, 2012). "Gun-toting Bale films in the area". Beaver County Times. Retrieved April 23, 2012.
Jump up ^ Tady, Scott (December 4, 2013). "Local sites appear in 'Out of the Furnace'". Beaver County Times. Retrieved December 4, 2013.
Jump up ^ "Academy Conversations: "Out of the Furnace"". Academy of Motion Picture Arts and Sciences. November 17, 2013. Retrieved November 30, 2013.
Jump up ^ "Three-Time Oscar®-Nominee Alberto Iglesias to Compose Original Score for Scortt Cooper's Currently-titled Out of the Furnace". Relativity Media. June 5, 2012. Retrieved July 10, 2013.
Jump up ^ "Dickon Hinchliffe Scoring ‘Hateship, Loveship’". Film Music Reporter. June 5, 2013. Retrieved July 14, 2013.
Jump up ^ "‘Out of the Furnace’ Soundtrack Details". Film Music Reporter. November 17, 2013. Retrieved November 17, 2013.
Jump up ^ Vancheri, Barbara (November 10, 2013). "Red-carpet footage from Out of the Furnace world premiere". Pittsburgh Post-Gazette. Retrieved November 30, 2013.
Jump up ^ "‘Out Of The Furnace’ Headed To LA & NYC Two Days Before Wide Release". Deadline.com. November 21, 2013. Retrieved November 30, 2013.
Jump up ^ Lyman, Eric J. (November 16, 2013). "Rome Film Fest: Italian Docudrama 'TIR' Wins Top Prize". The Hollywood Reporter. Retrieved November 30, 2013.
Jump up ^ Barton, Chris (December 7, 2013). "Box office: A chilly opening for 'Out of the Furnace'". Los Angeles Times. Retrieved December 7, 2013.
Jump up ^ Stewart, Andrew (December 8, 2013). "Box Office: 'Frozen' Ices 'Fire' in Weekend Showdown; 'Furnace' Fails to Gain Any Heat". Variety. Retrieved December 9, 2013.
^ Jump up to: a b Kaufman, Amy (December 8, 2013). "'Frozen' tops 'Catching Fire,' but 'Furnace' generates no heat". Los Angeles Times. Retrieved December 8, 2013.
Jump up ^ "Out of the Furnace (2013)". Rotten Tomatoes. Retrieved December 22, 2013.
Jump up ^ "Out of the Furnace". Metacritic. Retrieved December 13, 2013.
Jump up ^ http://www.metacritic.com/feature/film-critic-top-10-lists-best-movies-of-2013
Jump up ^ Duffy, Christie (December 4, 2013). "‘Out of the Furnace’ under fire in Mahwah". FiOS1. Retrieved December 5, 2013.
Jump up ^ Ivers, Dan (December 5, 2013). "Ramapough Indians, Mahwah officials decry depiction in new movie as 'hate crime'". NJ.com. Retrieved 2013-12-23.
Jump up ^ Incalcaterra, Laura (December 6, 2013). "Ramapough chief denounces Christian Bale movie as 'hate crime'". The Journal News. Retrieved December 7, 2013.
Jump up ^ Ma, Miles (December 23, 2013). "Members of Ramapough seek $50 million from makers of 'Out of the Furnace' A lawsuit says 'Out of the Furnace,' starring Christian Bale and Zoe Saldana, misrepresents members of the Ramapough Mountain Indians. (Relativity Media)". NJ.com. Retrieved 2013-12-23.
Jump up ^ "WILBUR C. DEGROAT, III, ARLITTA DEGROAT, LEON DEGROAT, DIANE DEGROAT, PIA DEGROAT DENNISON, MARIE DEGROAT MANN, RODNEY VAN DUNK, EUNICE DEGROAT, GILBERT DEGROAT,, ROSEMARIE MANN, AMANDA MANN, DAWN MANN, SANTANA PETERSON, JAZMINE PETERSON, RACHAEL MANN, DANIEL W. DENNISON, JR and JON VAN DUNK, Plaintiffs, v SCOTT COOPER; BRAD INGLESBY; RELATIVITY MEDIA, LLC a California limited liability company; APPIAN WAY, LLC, a California limited liability company; ENERGY ENTERTAINMENT, INC., a California corporation; SCOTT FREE PRODUCTIONS, INC., a California corporation; RED GRANITE PICTURES, INC. a California corporation; and JOHN DOES 1 to 5, Defendants" (PDF). NJ.com. Retrieved 2013-12-23.
Jump up ^ "Judge tosses out Ramapough Mountain Indians' lawsuit over Out of the Furnace". The Star Ledger. May 16, 2014. p. 17.
External links[edit]
Out of the Furnace on Facebook
Out of the Furnace on IMDb
Out of the Furnace at Rotten Tomatoes
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Crazy Heart (2009)
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Microevolution
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Evolutionary biology
Diagrammatic representation of the divergence of modern taxonomic groups from their common ancestor
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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
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External links[edit]
Microevolution (UC Berkeley)
Microevolution vs Macroevolution
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