text:
Call of Duty: WWII
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Call of Duty: WWII
Developer(s)
Sledgehammer Games
Publisher(s)
Activision
Director(s)
Bret Robbins
Dennis Adams
Artist(s)
Joe Salud
Composer(s)
Wilbert Roget II[1]
Series
Call of Duty
Platform(s)
Microsoft Windows
PlayStation 4
Xbox One
Release
WW: November 3, 2017
Genre(s)
First-person shooter
Mode(s)
Single-player, multiplayer
Call of Duty: WWII is an upcoming first-person shooter video game developed by Sledgehammer Games and published by Activision for Microsoft Windows, PlayStation 4 and Xbox One. It is the fourteenth primary installment and first Call of Duty series game to be set primarily during World War II since Call of Duty: World at War in 2008.[2] The game is set in European theatre of the war. The campaign is centered around a squad in the 1st Infantry Division, and follows their battles in the Western Front, while the multiplayer expands on different fronts not seen in the campaign. The game is scheduled to be released worldwide on November 3, 2017.
Contents
[hide]
1 Gameplay
1.1 Campaign
1.2 Multiplayer
1.3 Nazi Zombies
2 Plot
2.1 Campaign
2.2 Nazi Zombies
3 Development
4 References
5 External links
Gameplay[edit]
Similar to its predecessors, Call of Duty: WWII is a first-person shooter game but it removes the advanced system of movement present in the three previous Call of Duty titles, which included double jumping and wall running. Instead, it features a return of traditional movement to the series, taking it back to a "boots on the ground" gameplay style. The game does not feature an unlimited sprint mechanic, seen in the previous two titles.[3] Instead of a "slide" movement mechanic, which allowed players to slide quickly on the ground, WWII features a "hit-the-deck" mechanic that allows the player to leap forward and throw themselves on the ground in order to get to cover quickly, similarly to a previous mechanic known as "dolphin dive" in Treyarch's Call of Duty: Black Ops and Call of Duty: Black Ops II.[4]
Campaign[edit]
WWII is the first title since the original Call of Duty to not feature health regeneration in the campaign. Instead, players must rely on their medic squadmates to provide health packs, as well as other squad members to provide ammunition. Other squad members can spot enemies, revealing their position in form of silhouettes.[5] Enemy soldiers in the campaign can be captured, and wounded allies can be dragged to cover. Both of these actions, as well as killing enemies and completing objectives, add up to player's experience (XP) which can be increased and improve the character's skills. In some parts of the campaign, players will be able to control vehicles.[6]
Multiplayer[edit]
The multiplayer mode for Call of Duty: WWII was revealed on E3 2017, which took place from June 13–15.[7] Sledgehammer Games has announced new features such as the new headquarters social space, divisions, war mode and the return to "boots-on-the-ground" gameplay. Players who pre-order the game were invited to a closed beta, which was released initially for the PlayStation 4, but later was released on other platforms.[8]
In online multiplayer matches, players are randomly assigned either to Allies or Axis side. On Allies side, players can play as soldiers in American, British, or French Resistance armies. On the Axis side, players play directly as Germans. When explaining which German Axis powers would be playable, Glen Schofield, co-founder and co-studio head at Sledgehammer, appeared to state that the Clean Wehrmacht myth was the criteria they had used in their decision, saying: "The big distinction that Germans still make today is that between the German military and the Nazis. We made sure we made that distinction in the game, that the Germans were doing their duty.”[9]
Instead of the usual create-a-class system, WWII features Divisions. Players can choose one out of five divisions, each with their own different basic combat training, division training and weapon skills. This also eliminates perks, as players need to progress through ranks in divisions in order to use additional skills. Five divisions featured in the game are:
Infantry: The most versatile division in the game, it is equipped for mid to long range combat. The special skill of this division is the bayonet charge, which can lead to brutal melee combat.
Airborne: This division makes players move fast while remaining quiet. Players in this division can attach suppressors to submachine guns at any time during combat which allows for more stealth based gameplay style.
Armored: This division has the heaviest firepower, being able to mount machine guns and carry rocket launchers as secondary weapons.
Mountain: This division focuses on long range combat, making sniper rifles more precise with aim assist and improving the aim through scope by blocking out surroundings from player's view.
Expeditionary: Shotguns used by players in this division have incendiary rounds that burn enemies to death.
WWII also features Headquarters mode, which acts as a social space in the game. The massive hub is set on the Omaha Beach in Normandy, three days after the invasion when Allies retake the beach and turn it into a base. 48 players can be in the Headquarters at a time, and take part in various activities. For example, players can watch other players open loot boxes while in the Headquarters.[10] There is a firing range in the hub, where all players can practise their shooting skills with all weapons, as well as a field where they can test scorestreaks. There are also areas where players can engage in "1v1" fights, as other players watch the duels.[11]
The end-game "killcam" highlight shown at the end of every Multiplayer match has been changed to "Bronze Star", which only show kills "deemed most impressive," similar to Overwatch.[12] The only exception to this is the Search & Destroy game mode, which does not use Bronze Star killcam.[13]
Nazi Zombies[edit]
WWII will include a zombies cooperative game mode, similar to previous entries by Treyarch and Infinity Ward, with its own original storyline set in alternate history and separate from the campaign. The game mode, dubbed Nazi Zombies in reference to its first iteration in Treyarch's Call of Duty: World at War, is also set in the events of World War II, as the Third Reich makes a desperate attempt to turn the tide of the war by creating an undead army in the final stages of the war. While the mode is based on science-fiction and is a fictional take on the war, Sledgehammer Games co-founder Michael Condrey said that the story of the mode is based on some "real events". He also revealed that the experience is similar to Dead Space, a third person shooter horror video game directed by both Condrey and Schofield during their work time at EA Redwood Shores.[14]
In regards to gameplay, Nazi Zombies retains the wave-based survival formula that have been used in all previous Zombies entries, with brand new additions. A class system is introduced, where players can opt for one of four combat roles: Offense, Control, Medic and Support, which provide different in-game abilities. Class loadouts are also included, with equippable perks similar to the multiplayer mode. Sledgehammer Games also attempted to rationalize some of the popular mechanics, such as weapon wallbuys and currencies, with realistic explanation that fit within the lore of the game mode.
Plot[edit]
Campaign[edit]
Call of Duty: WWII focuses on the actions of a 12-men squad of the American 1st Infantry Division from 1944 to 1945 in the European theater of conflict, at a time when the Allied forces were starting to gather strength on their march into Germany. The campaign will cover fighting in occupied France and Belgium, and across the Rhine river valley into the heart of Germany.[7] Some of the iconic historical battles covered in the campaign will be the Invasion of Normandy (D-Day), Liberation of Paris, Battle of Aachen, Battle of Hürtgen Forest and Battle of the Bulge. Although the story is primarily set in 1944 and 1945, some parts of it will "touch on" events that happened from 1940 to 1944 to "help set the stage for the narrative and establish the characters."[15]
In the single-player campaign, players primarily play as two soldiers of the 16th Infantry Regiment: Private Ronald "Red" Daniels (Brett Zimmerman) and his best friend, Jewish American, Private First Class Robert Zussman (Jonathan Tucker). Additional playable characters include Rousseau, a female Maquis leader of the French Forces of the Interior, and Major Arthur Crowley, a British Special Operations Executive officer a RAF Pilot and Tank Commander. Daniels' and Zussman's platoon mates include Technical Sergeant William Pierson (Josh Duhamel), and platoon leader, 1st Lieutenant Joseph Turner (Jeffrey Pierce).[16] Pittsburgh Steelers running back Le'Veon Bell and offensive tackle Alejandro Villanueva are set to make a cameo appearance in the game.[17]
Nazi Zombies[edit]
The Nazi Zombies story of WWII takes place in an alternate history, where the Nazis have conducted research on necromancy based on the legend of King Frederick Barbarossa's hidden treasure. The story is centered around Marie Fischer (Katheryn Winnick), a young German engineer working with the Office of Strategic Services. Marie's brother, Klaus, has taken part in the research of Barbarossa's treasure, working with Doctor Peter Straub (Udo Kier). Marie is joined by three companions: ex-art thief Drostan Hynd (David Tennant), ex-art historian and French Resistance fighter Olivia Durant (Élodie Yung), and military sergeant Jefferson Potts (Ving Rhames), as they attempt to conduct a covert operation to learn of the Nazis' plans, retrieve the stolen artifacts and rescue Klaus in the process.
Development[edit]
A modern Call of Duty title set in World War II was alluded to in a 2014 Call of Duty: Advanced Warfare launch interview with Michael Condrey, co-founder of Sledgehammer Games. In the interview by Metro, the interviewer asked him what the possibilities of where the next Call of Duty could go in terms of setting. Condrey responded, "Well, no. It’s curious. I can only answer from my own personal tastes, this is my own personal opinion. But some of my favourite pieces of entertainment are set in World War II. Band of Brothers, I’m a massive fan of Band of Brothers." Condrey then dived further into the subject, "And that’s a great hero’s war, kind of the last that was recognised as a noble cause in a war. So yeah, I think a next generation game with the latest production values and robustness in a World War II setting like Band of Brothers would be amazing. Now, how would it play and how would the multiplayer work after the new movement set in Advanced Warfare? That’s a tougher question than I’ve had to tackle yet…".[18]
Sledgehammer Games have yet to reveal all the authentic settings from World War II that developers have put into the game's storyline. Activision is refusing to deny claims that Nazi extermination camps could feature in the game.[9] Adam Rosenberg of Mashable wrote that video games set during World War II tended to be "Holocaust deniers" in the sense that they avoided broaching the subject for business reasons, but that this could be the very first Call of Duty World War II based game where the Holocaust would be depicted. Senior creative director Bret Robbins said in an interview "Some very, very dark things happened during this conflict and it felt wrong for us to ignore that." He further stated “We absolutely show atrocities. It’s an unfortunate part of the history, but you can’t tell an authentic, truthful story without going there. So we went there.” Robbins argued that audiences can now handle games with more maturity and nuance, “People are ready for it. They want it,” he said.[19][20] When asked directly over Twitter as to whether or not the story campaign would allow gamers the opportunity to play as soldiers from the Axis powers such as Nazi Germany and Fascist Italy, Sledgehammer Games confirmed that the campaign gameplay would be limited to allied forces. More specifically, Sledgehammer co-founder Michael Condrey confirmed that the game will focus exclusively on the Allied powers.[9]
The Windows version of the game was developed in partnership with Raven Software. In regards to it, Raven's CTO Dwight Luetscher stated that they were trying to focus on the Windows platform, as well as the community, by responding to their needs for it to excel. The Windows version features several notable changes, including removal of controller aim assist and addition of sensitivity slider for aim-down-sights mechanic.[21]
All pre-orders excluding the PC version include access to the private beta, which was made available first on the PlayStation 4 from August 25 to 28, followed by a second week for both PlayStation 4 and Xbox One from September 1 to 4. The PC beta was announced as an open beta, and ran from September 29 to October 2 on Steam. Players who participated in the beta receive the Beta Combat Pack for the full game, which contains a special in-game helmet, emblem and calling card. The game will be available in 3 editions: Base Edition, Digital Deluxe Edition, and the Pro Edition. The Pro Edition is sold exclusively at GameStop. Any pre-orders of the game at GameStop also included a limited edition hat.[22] The game will be released worldwide on November 3, 2017 for Microsoft Windows, PlayStation 4 and Xbox One.[23]
References[edit]
Jump up ^ McMillan, Emily. "Wilbert Roget II composes for Call of Duty: World War II". Video Game Music Online. Retrieved 20 October 2017.
Jump up ^ Campbell, Colin (April 21, 2017). "Call of Duty: WWII confirmed, full reveal next week". Polygon. Retrieved April 26, 2017.
Jump up ^ "Unlimited sprint won’t be featured in Call of Duty: WWII". Retrieved July 11, 2017.
Jump up ^ Condrey, Michael (June 13, 2017). "You can "hit-the-deck" and scramble for cover, officially.". @MichaelCondrey. Retrieved July 11, 2017.
Jump up ^ Dayus, Oscar (April 27, 2017). "Call Of Duty: WW2 Ditching Health Regen, Has Some Form Of Dedicated Classes". GameSpot. Retrieved June 16, 2017.
Jump up ^ acast (June 16, 2017). "E3 2017: Call Of Duty: WW2 Interview | Gamertag Radio on acast". acast. Retrieved July 12, 2017.
^ Jump up to: a b Kelly, Kevin (April 26, 2017). "Announcing Call of Duty: WWII". Activision. Retrieved April 27, 2017.
Jump up ^ Kain, Erik (April 26, 2017). "Everything You Need To Know About The 'Call Of Duty: World War II' Reveal". Forbes. Retrieved June 16, 2017.
^ Jump up to: a b c Wright, James (May 28, 2017). "Call of Duty WW2: Bosses reveal the one gameplay feature they WON'T include". Daily Star. Retrieved June 16, 2017.
Jump up ^ "Call of Duty: WW2 Gives You Prizes For Watching Others Open Loot Boxes". Kotaku UK. Retrieved August 10, 2017.
Jump up ^ Stuart, Keith (June 28, 2017). "Call of Duty: WWII – how an indie classic inspired the latest instalment". The Guardian. ISSN 0261-3077. Retrieved July 12, 2017.
Jump up ^ "Call of Duty: WWII Ditches Endgame Killcam for Something Better". twinfinite.net. Retrieved August 25, 2017.
Jump up ^ Condrey, Michael. "I'm way overdue on this, sorry. Yes, S&D will be final killcam, not Bronze Star camera.". Twitter. Retrieved September 12, 2017.
Jump up ^ Keshav (June 3, 2017). "Call of Duty: WWII Zombies has a 'back story that’s based on real events,' similar experience to Dead Space". Charlie INTEL. Retrieved July 12, 2017.
Jump up ^ Dayus, Oscar (April 27, 2017). "Call Of Duty: WW2 Ditching Health Regen, Has Some Form Of Dedicated Classes". GameSpot. Retrieved July 12, 2017.
Jump up ^ Hurley, Leon (April 26, 2017). "Call of Duty WW2 will feature a playable French Resistance woman, and "a global cast" that includes a German family". Games Radar. Retrieved June 16, 2017.
Jump up ^ Gentille, Sean (September 19, 2017). "'Call of Duty: WWII' features Le'Veon Bell and Alejandro Villanueva. Here's what they think about that.". Pittsburgh Post-Gazette. Retrieved September 19, 2017.
Jump up ^ Metro.co.uk, David Jenkins for (November 3, 2014). "COD: Advanced Warfare Michael Condrey interview". Metro. Retrieved July 13, 2017.
Jump up ^ Rosenberg, Adam (April 26, 2017). "'Call of Duty: WWII' won't ignore the Holocaust anymore". Mashable. Retrieved June 16, 2017.
Jump up ^ Pink, Aiden (May 3, 2017). "New ‘Call Of Duty’ Video Game Will Depict The Holocaust". The Forward. Retrieved June 16, 2017.
Jump up ^ Fenlon, Wes (September 28, 2017). "Call of Duty: WWII devs want to 'win the hearts and minds of the PC community'". PC Gamer. Retrieved September 28, 2017.
Jump up ^ Plagge, Kallie (April 26, 2017). "Call Of Duty: WW2 - Everything We Know So Far". GameSpot. Retrieved June 16, 2017.
Jump up ^ Webster, Andrew (April 26, 2017). "Call of Duty: WWII is launching November 3rd". The Verge. Retrieved April 26, 2017.
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Karyotype
From Wikipedia, the free encyclopedia
Jump to: navigation, search
"Cytotype" redirects here. For other uses, see Cytotype (disambiguation).
A karyotype is the number and appearance of chromosomes in the nucleus of a eukaryotic cell. The term is also used for the complete set of chromosomes in a species or in an individual organism[1][2][3] and for a test that detects this complement or measures the number.
Karyotypes describe the chromosome count of an organism and what these chromosomes look like under a light microscope. Attention is paid to their length, the position of the centromeres, banding pattern, any differences between the sex chromosomes, and any other physical characteristics.[4] The preparation and study of karyotypes is part of cytogenetics.
Karyogram of human male using Giemsa staining
The study of whole sets of chromosomes is sometimes known as karyology. The chromosomes are depicted (by rearranging a photomicrograph) in a standard format known as a karyogram or idiogram: in pairs, ordered by size and position of centromere for chromosomes of the same size.
The basic number of chromosomes in the somatic cells of an individual or a species is called the somatic number and is designated 2n. In the germ-line (the sex cells) the chromosome number is n (humans: n = 23).[2]p28 Thus, in humans 2n = 46.
So, in normal diploid organisms, autosomal chromosomes are present in two copies. There may, or may not, be sex chromosomes. Polyploid cells have multiple copies of chromosomes and haploid cells have single copies.
The study of karyotypes is important for cell biology and genetics, and the results may be used in evolutionary biology (karyosystematics)[5] and medicine. Karyotypes can be used for many purposes; such as to study chromosomal aberrations, cellular function, taxonomic relationships, and to gather information about past evolutionary events.
Contents
[hide]
1 History of karyotype studies
2 Observations on karyotypes
2.1 Staining
2.2 Observations
2.3 Human karyotype
3 Diversity and evolution of karyotypes
3.1 Changes during development
3.2 Number of chromosomes in a set
3.3 Fundamental number
3.4 Ploidy
3.5 Aneuploidy
3.6 Chromosomal polymorphism
3.7 Species trees
3.8 Chromosome banding
4 Depiction of karyotypes
4.1 Types of banding
4.2 Classic karyotype cytogenetics
4.3 Multicolor FISH (mFISH) and spectral karyotype (SKY technique)
4.4 Digital karyotyping
5 Chromosome abnormalities
6 See also
7 References
8 External links
History of karyotype studies[edit]
Chromosomes were first observed in plant cells by Carl Wilhelm von Nägeli in 1842. Their behavior in animal (salamander) cells was described by Walther Flemming, the discoverer of mitosis, in 1882. The name was coined by another German anatomist, Heinrich von Waldeyer in 1888. It is New Latin from Ancient Greek κάρυον karyon, "kernel", "seed", or "nucleus", and τύπος typos, "general form").
The next stage took place after the development of genetics in the early 20th century, when it was appreciated that chromosomes (that can be observed by karyotype) were the carrier of genes. Lev Delaunay (ru) seems to have been the first person to define the karyotype as the phenotypic appearance of the somatic chromosomes, in contrast to their genic contents (1922).[6][7] The subsequent history of the concept can be followed in the works of C. D. Darlington[8] and Michael JD White.[2][9]
Investigation into the human karyotype took many years to settle the most basic question: how many chromosomes does a normal diploid human cell contain?[10] In 1912, Hans von Winiwarter reported 47 chromosomes in spermatogonia and 48 in oogonia, concluding an XX/XO sex determination mechanism.[11] Painter in 1922 was not certain whether the diploid of humans was 46 or 48, at first favoring 46,[12] but revised his opinion from 46 to 48, and he correctly insisted on humans having an XX/XY system.[13] Considering the techniques of the time, these results were remarkable.
Fusion of ancestral chromosomes left distinctive remnants of telomeres, and a vestigial centromere
In textbooks, the number of human chromosomes remained at 48 for over thirty years. New techniques were needed to correct this error. Joe Hin Tjio working in Albert Levan's lab[14] was responsible for finding the approach:
Using cells in tissue culture
Pretreating cells in a hypotonic solution, which swells them and spreads the chromosomes
Arresting mitosis in metaphase by a solution of colchicine
Squashing the preparation on the slide forcing the chromosomes into a single plane
Cutting up a photomicrograph and arranging the result into an indisputable karyogram.
The work took place in 1955, and was published in 1956. The karyotype of humans includes only 46 chromosomes.[15][16] Rather interestingly, the great apes have 48 chromosomes. Human chromosome 2 is now known to be a result of an end-to-end fusion of two ancestral ape chromosomes.[17][18]
Observations on karyotypes[edit]
Staining[edit]
The study of karyotypes is made possible by staining. Usually, a suitable dye, such as Giemsa,[19] is applied after cells have been arrested during cell division by a solution of colchicine usually in metaphase or prometaphase when most condensed. In order for the Giemsa stain to adhere correctly, all chromosomal proteins must be digested and removed. For humans, white blood cells are used most frequently because they are easily induced to divide and grow in tissue culture.[20] Sometimes observations may be made on non-dividing (interphase) cells. The sex of an unborn fetus can be determined by observation of interphase cells (see amniotic centesis and Barr body).
Observations[edit]
Six different characteristics of karyotypes are usually observed and compared:[21]
Differences in absolute sizes of chromosomes. Chromosomes can vary in absolute size by as much as twenty-fold between genera of the same family. For example, the legumes Lotus tenuis and Vicia faba each have six pairs of chromosomes, yet V. faba chromosomes are many times larger. These differences probably reflect different amounts of DNA duplication.
Differences in the position of centromeres. These differences probably came about through translocations.
Differences in relative size of chromosomes. These differences probably arose from segmental interchange of unequal lengths.
Differences in basic number of chromosomes. These differences could have resulted from successive unequal translocations which removed all the essential genetic material from a chromosome, permitting its loss without penalty to the organism (the dislocation hypothesis) or through fusion. Humans have one pair fewer chromosomes than the great apes. Human chromosome 2 appears to have resulted from the fusion of two ancestral chromosomes, and many of the genes of those two original chromosomes have been translocated to other chromosomes.
Differences in number and position of satellites. Satellites are small bodies attached to a chromosome by a thin thread.
Differences in degree and distribution of heterochromatic regions. Heterochromatin stains darker than euchromatin. Heterochromatin is packed tighter. Heterochromatin consists mainly of genetically inactive and repetitive DNA sequences as well as containing a larger amount of Adenine-Thymine pairs. Euchromatin is usually under active transcription and stains much lighter as it has less affinity for the giemsa stain.[22] Euchromatin regions contain larger amounts of Guanine-Cytosine pairs. The staining technique using giemsa staining is called G banding and therefore produces the typical "G-Bands".[22]
A full account of a karyotype may therefore include the number, type, shape and banding of the chromosomes, as well as other cytogenetic information.
Variation is often found:
between the sexes,
between the germ-line and soma (between gametes and the rest of the body),
between members of a population (chromosome polymorphism),
in geographic specialization, and
in mosaics or otherwise abnormal individuals.[9]
Human karyotype[edit]
human karyotype
The normal human karyotypes contain 22 pairs of autosomal chromosomes and one pair of sex chromosomes (allosomes). Normal karyotypes for females contain two X chromosomes and are denoted 46,XX; males have both an X and a Y chromosome denoted 46,XY. Any variation from the standard karyotype may lead to developmental abnormalities.
Diversity and evolution of karyotypes[edit]
Although the replication and transcription of DNA is highly standardized in eukaryotes, the same cannot be said for their karyotypes, which are highly variable. There is variation between species in chromosome number, and in detailed organization, despite their construction from the same macromolecules. This variation provides the basis for a range of studies in evolutionary cytology.
In some cases there is even significant variation within species. In a review, Godfrey and Masters conclude:
In our view, it is unlikely that one process or the other can independently account for the wide range of karyotype structures that are observed ... But, used in conjunction with other phylogenetic data, karyotypic fissioning may help to explain dramatic differences in diploid numbers between closely related species, which were previously inexplicable.[23]
Although much is known about karyotypes at the descriptive level, and it is clear that changes in karyotype organization has had effects on the evolutionary course of many species, it is quite unclear what the general significance might be.
We have a very poor understanding of the causes of karyotype evolution, despite many careful investigations ... the general significance of karyotype evolution is obscure.
—  Maynard Smith[24]
Changes during development[edit]
Instead of the usual gene repression, some organisms go in for large-scale elimination of heterochromatin, or other kinds of visible adjustment to the karyotype.
Chromosome elimination. In some species, as in many sciarid flies, entire chromosomes are eliminated during development.[25]
Chromatin diminution (founding father: Theodor Boveri). In this process, found in some copepods and roundworms such as Ascaris suum, portions of the chromosomes are cast away in particular cells. This process is a carefully organised genome rearrangement where new telomeres are constructed and certain heterochromatin regions are lost.[26][27] In A. suum, all the somatic cell precursors undergo chromatin diminution.[28]
X-inactivation. The inactivation of one X chromosome takes place during the early development of mammals (see Barr body and dosage compensation). In placental mammals, the inactivation is random as between the two Xs; thus the mammalian female is a mosaic in respect of her X chromosomes. In marsupials it is always the paternal X which is inactivated. In human females some 15% of somatic cells escape inactivation,[29] and the number of genes affected on the inactivated X chromosome varies between cells: in fibroblast cells up about 25% of genes on the Barr body escape inactivation.[30]
Number of chromosomes in a set[edit]
A spectacular example of variability between closely related species is the muntjac, which was investigated by Kurt Benirschke and his colleague Doris Wurster. The diploid number of the Chinese muntjac, Muntiacus reevesi, was found to be 46, all telocentric. When they looked at the karyotype of the closely related Indian muntjac, Muntiacus muntjak, they were astonished to find it had female = 6, male = 7 chromosomes.[31]
They simply could not believe what they saw ... They kept quiet for two or three years because they thought something was wrong with their tissue culture ... But when they obtained a couple more specimens they confirmed [their findings].
—  Hsu p. 73-4[16]
The number of chromosomes in the karyotype between (relatively) unrelated species is hugely variable. The low record is held by the nematode Parascaris univalens, where the haploid n = 1; and an ant: Myrmecia pilosula.[32] The high record would be somewhere amongst the ferns, with the adder's tongue fern Ophioglossum ahead with an average of 1262 chromosomes.[33] Top score for animals might be the shortnose sturgeon Acipenser brevirostrum at 372 chromosomes.[34] The existence of supernumerary or B chromosomes means that chromosome number can vary even within one interbreeding population; and aneuploids are another example, though in this case they would not be regarded as normal members of the population.
Fundamental number[edit]
The fundamental number, FN, of a karyotype is the number of visible major chromosomal arms per set of chromosomes.[35][36] Thus, FN ≤ 2 x 2n, the difference depending on the number of chromosomes considered single-armed (acrocentric or telocentric) present. Humans have FN = 82,[37] due to the presence of five acrocentric chromosome pairs: 13, 14, 15, 21, and 22 (the human Y chromosome is also acrocentric). The fundamental autosomal number or autosomal fundamental number, FNa[38] or AN,[39] of a karyotype is the number of visible major chromosomal arms per set of autosomes (non-sex-linked chromosomes).
Ploidy[edit]
Ploidy is the number of complete sets of chromosomes in a cell.
Polyploidy, where there are more than two sets of homologous chromosomes in the cells, occurs mainly in plants. It has been of major significance in plant evolution according to Stebbins.[40][41][42][43] The proportion of flowering plants which are polyploid was estimated by Stebbins to be 30–35%, but in grasses the average is much higher, about 70%.[44] Polyploidy in lower plants (ferns, horsetails and psilotales) is also common, and some species of ferns have reached levels of polyploidy far in excess of the highest levels known in flowering plants.
Polyploidy in animals is much less common, but it has been significant in some groups.[45]
Polyploid series in related species which consist entirely of multiples of a single basic number are known as euploid.
Haplo-diploidy, where one sex is diploid, and the other haploid. It is a common arrangement in the Hymenoptera, and in some other groups.
Endopolyploidy occurs when in adult differentiated tissues the cells have ceased to divide by mitosis, but the nuclei contain more than the original somatic number of chromosomes.[46] In the endocycle (endomitosis or endoreduplication) chromosomes in a 'resting' nucleus undergo reduplication, the daughter chromosomes separating from each other inside an intact nuclear membrane.[47]
In many instances, endopolyploid nuclei contain tens of thousands of chromosomes (which cannot be exactly counted). The cells do not always contain exact multiples (powers of two), which is why the simple definition 'an increase in the number of chromosome sets caused by replication without cell division' is not quite accurate.
This process (especially studied in insects and some higher plants such as maize) may be a developmental strategy for increasing the productivity of tissues which are highly active in biosynthesis.[48]
The phenomenon occurs sporadically throughout the eukaryote kingdom from protozoa to humans; it is diverse and complex, and serves differentiation and morphogenesis in many ways.[49]
See palaeopolyploidy for the investigation of ancient karyotype duplications.
Aneuploidy[edit]
Aneuploidy is the condition in which the chromosome number in the cells is not the typical number for the species. This would give rise to a chromosome abnormality such as an extra chromosome or one or more chromosomes lost. Abnormalities in chromosome number usually cause a defect in development. Down syndrome and Turner syndrome are examples of this.
Aneuploidy may also occur within a group of closely related species. Classic examples in plants are the genus Crepis, where the gametic (= haploid) numbers form the series x = 3, 4, 5, 6, and 7; and Crocus, where every number from x = 3 to x = 15 is represented by at least one species. Evidence of various kinds shows that trends of evolution have gone in different directions in different groups.[50] Closer to home, the great apes have 24x2 chromosomes whereas humans have 23x2. Human chromosome 2 was formed by a merger of ancestral chromosomes, reducing the number.[51]
Chromosomal polymorphism[edit]
Some species are polymorphic for different chromosome structural forms.[52] The structural variation may be associated with different numbers of chromosomes in different individuals, which occurs in the ladybird beetle Chilocorus stigma, some mantids of the genus Ameles,[citation needed] the European shrew Sorex araneus.[53] There is some evidence from the case of the mollusc Thais lapillus (the dog whelk) on the Brittany coast, that the two chromosome morphs are adapted to different habitats.[54]
Species trees[edit]
The detailed study of chromosome banding in insects with polytene chromosomes can reveal relationships between closely related species: the classic example is the study of chromosome banding in Hawaiian drosophilids by Hampton L. Carson.
In about 6,500 sq mi (17,000 km2), the Hawaiian Islands have the most diverse collection of drosophilid flies in the world, living from rainforests to subalpine meadows. These roughly 800 Hawaiian drosophilid species are usually assigned to two genera, Drosophila and Scaptomyza, in the family Drosophilidae.
The polytene banding of the 'picture wing' group, the best-studied group of Hawaiian drosophilids, enabled Carson to work out the evolutionary tree long before genome analysis was practicable. In a sense, gene arrangements are visible in the banding patterns of each chromosome. Chromosome rearrangements, especially inversions, make it possible to see which species are closely related.
The results are clear. The inversions, when plotted in tree form (and independent of all other information), show a clear "flow" of species from older to newer islands. There are also cases of colonization back to older islands, and skipping of islands, but these are much less frequent. Using K-Ar dating, the present islands date from 0.4 million years ago (mya) (Mauna Kea) to 10mya (Necker). The oldest member of the Hawaiian archipelago still above the sea is Kure Atoll, which can be dated to 30 mya. The archipelago itself (produced by the Pacific plate moving over a hot spot) has existed for far longer, at least into the Cretaceous. Previous islands now beneath the sea (guyots) form the Emperor Seamount Chain.[55]
All of the native Drosophila and Scaptomyza species in Hawaiʻi have apparently descended from a single ancestral species that colonized the islands, probably 20 million years ago. The subsequent adaptive radiation was spurred by a lack of competition and a wide variety of niches. Although it would be possible for a single gravid female to colonise an island, it is more likely to have been a group from the same species.[56][57][58][59]
There are other animals and plants on the Hawaiian archipelago which have undergone similar, if less spectacular, adaptive radiations.[60][61]
Chromosome banding[edit]
Chromosomes display a banded pattern when treated with some stains. Bands are alternating light and dark stripes that appear along the lengths of chromosomes. Unique banding patterns are used to identify chromosomes and to diagnose chromosomal aberrations, including chromosome breakage, loss, duplication, translocation or inverted segments. A range of different chromosome treatments produce a range of banding patterns: G-bands, R-bands, C-bands, Q-bands, T-bands and NOR-bands.
Depiction of karyotypes[edit]
Types of banding[edit]
Cytogenetics employs several techniques to visualize different aspects of chromosomes:[20]
G-banding is obtained with Giemsa stain following digestion of chromosomes with trypsin. It yields a series of lightly and darkly stained bands — the dark regions tend to be heterochromatic, late-replicating and AT rich. The light regions tend to be euchromatic, early-replicating and GC rich. This method will normally produce 300–400 bands in a normal, human genome.
R-banding is the reverse of G-banding (the R stands for "reverse"). The dark regions are euchromatic (guanine-cytosine rich regions) and the bright regions are heterochromatic (thymine-adenine rich regions).
C-banding: Giemsa binds to constitutive heterochromatin, so it stains centromeres.The name is derived from centromeric or constitutive heterochromatin. The preparations undergo alkaline denaturation prior to staining leading to an almost complete depurination of the DNA. After washing the probe the remaining DNA is renatured again and stained with Giemsa solution consisting of methylene azure, methylene violet, methylene blue, and eosin. Heterochromatin binds a lot of the dye, while the rest of the chromosomes absorb only little of it. The C-bonding proved to be especially well-suited for the characterization of plant chromosomes.
Q-banding is a fluorescent pattern obtained using quinacrine for staining. The pattern of bands is very similar to that seen in G-banding.They can be recognized by a yellow fluorescence of differing intensity. Most part of the stained DNA is heterochromatin. Quinacrin (atebrin) binds both regions rich in AT and in GC, but only the AT-quinacrin-complex fluoresces. Since regions rich in AT are more common in heterochromatin than in euchromatin, these regions are labelled preferentially. The different intensities of the single bands mirror the different contents of AT. Other fluorochromes like DAPI or Hoechst 33258 lead also to characteristic, reproducible patterns. Each of them produces its specific pattern. In other words: the properties of the bonds and the specificity of the fluorochromes are not exclusively based on their affinity to regions rich in AT. Rather, the distribution of AT and the association of AT with other molecules like histones, for example, influences the binding properties of the fluorochromes.
T-banding: visualize telomeres.
Silver staining: Silver nitrate stains the nucleolar organization region-associated protein. This yields a dark region where the silver is deposited, denoting the activity of rRNA genes within the NOR.
Classic karyotype cytogenetics[edit]
Karyogram from a human female lymphocyte probed for the Alu sequence using FISH.
In the "classic" (depicted) karyotype, a dye, often Giemsa (G-banding), less frequently mepacrine (quinacrine), is used to stain bands on the chromosomes. Giemsa is specific for the phosphate groups of DNA. Quinacrine binds to the adenine-thymine-rich regions. Each chromosome has a characteristic banding pattern that helps to identify them; both chromosomes in a pair will have the same banding pattern.
Karyotypes are arranged with the short arm of the chromosome on top, and the long arm on the bottom. Some karyotypes call the short and long arms p and q, respectively. In addition, the differently stained regions and sub-regions are given numerical designations from proximal to distal on the chromosome arms. For example, Cri du chat syndrome involves a deletion on the short arm of chromosome 5. It is written as 46,XX,5p-. The critical region for this syndrome is deletion of p15.2 (the locus on the chromosome), which is written as 46,XX,del(5)(p15.2).[62]
Multicolor FISH (mFISH) and spectral karyotype (SKY technique)[edit]
Spectral karyogram of a human female
Multicolor FISH and the older spectral karyotyping are molecular cytogenetic techniques used to simultaneously visualize all the pairs of chromosomes in an organism in different colors. Fluorescently labeled probes for each chromosome are made by labeling chromosome-specific DNA with different fluorophores. Because there are a limited number of spectrally distinct fluorophores, a combinatorial labeling method is used to generate many different colors. Fluorophore combinations are captured and analyzed by a fluorescence microscope using up to 7 narrow-banded fluorescence filters or, in the case of spectral karyotyping, by using an interferometer attached to a fluorescence microscope. In the case of an mFISH image, every combination of fluorochromes from the resulting original images is replaced by a pseudo color in a dedicated image analysis software. Thus, chromosomes or chromosome sections can be visualized and identified, allowing for the analysis of chromosomal rearrangements.[63] In the case of spectral karyotyping, image processing software assigns a pseudo color to each spectrally different combination, allowing the visualization of the individually colored chromosomes.[64]
Spectral human karyotype
Multicolor FISH is used to identify structural chromosome aberrations in cancer cells and other disease conditions when Giemsa banding or other techniques are not accurate enough.
Digital karyotyping[edit]
Digital karyotyping is a technique used to quantify the DNA copy number on a genomic scale. Short sequences of DNA from specific loci all over the genome are isolated and enumerated.[65] This method is also known as virtual karyotyping.
Chromosome abnormalities[edit]
Main article: Chromosome abnormalities
Chromosome abnormalities can be numerical, as in the presence of extra or missing chromosomes, or structural, as in derivative chromosome, translocations, inversions, large-scale deletions or duplications. Numerical abnormalities, also known as aneuploidy, often occur as a result of nondisjunction during meiosis in the formation of a gamete; trisomies, in which three copies of a chromosome are present instead of the usual two, are common numerical abnormalities. Structural abnormalities often arise from errors in homologous recombination. Both types of abnormalities can occur in gametes and therefore will be present in all cells of an affected person's body, or they can occur during mitosis and give rise to a genetic mosaic individual who has some normal and some abnormal cells.
Chromosomal abnormalities that lead to disease in humans include
Turner syndrome results from a single X chromosome (45,X or 45,X0).
Klinefelter syndrome, the most common male chromosomal disease, otherwise known as 47,XXY, is caused by an extra X chromosome.
Edwards syndrome is caused by trisomy (three copies) of chromosome 18.
Down syndrome, a common chromosomal disease, is caused by trisomy of chromosome 21.
Patau syndrome is caused by trisomy of chromosome 13.
Trisomy 9, believed to be the 4th most common trisomy, has many long lived affected individuals but only in a form other than a full trisomy, such as trisomy 9p syndrome or mosaic trisomy 9. They often function quite well, but tend to have trouble with speech.
Also documented are trisomy 8 and trisomy 16, although they generally do not survive to birth.
Some disorders arise from loss of just a piece of one chromosome, including
Cri du chat (cry of the cat), from a truncated short arm on chromosome 5. The name comes from the babies' distinctive cry, caused by abnormal formation of the larynx.
1p36 Deletion syndrome, from the loss of part of the short arm of chromosome 1.
Angelman syndrome – 50% of cases have a segment of the long arm of chromosome 15 missing; a deletion of the maternal genes, example of imprinting disorder.
Prader-Willi syndrome – 50% of cases have a segment of the long arm of chromosome 15 missing; a deletion of the paternal genes, example of imprinting disorder.
Chromosomal abnormalities can also occur in cancerous cells of an otherwise genetically normal individual; one well-documented example is the Philadelphia chromosome, a translocation mutation commonly associated with chronic myelogenous leukemia and less often with acute lymphoblastic leukemia.
See also[edit]
Cytogenetic notation
Genome screen
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Jump up ^ Lisa G. Shaffer; Niels Tommerup, eds. (2005). ISCN 2005: An International System for Human Cytogenetic Nomenclature. Switzerland: S. Karger AG. ISBN 3-8055-8019-3.
Jump up ^ Liehr T, Starke H, Weise A, Lehrer H, Claussen U (January 2004). "Multicolour FISH probe sets and their applications". Histol Histopathol. 19 (1): 229–237. PMID 14702191.
Jump up ^ Schröck E, du Manoir S, Veldman T, et al. (July 1996). "Multicolor spectral karyotyping of human chromosomes". Science. 273 (5274): 494–7. Bibcode:1996Sci...273..494S. doi:10.1126/science.273.5274.494. PMID 8662537.
Jump up ^ Wang TL, Maierhofer C, Speicher MR, et al. (December 2002). "Digital karyotyping". Proc. Natl. Acad. Sci. U.S.A. 99 (25): 16156–61. Bibcode:2002PNAS...9916156W. doi:10.1073/pnas.202610899. PMC 138581  . PMID 12461184.
External links[edit]
Media related to Karyotypes at Wikimedia Commons
Making a karyotype, an online activity from the University of Utah's Genetic Science Learning Center.
Karyotyping activity with case histories from the University of Arizona's Biology Project.
Printable karyotype project from Biology Corner, a resource site for biology and science teachers.
Chromosome Staining and Banding Techniques
Bjorn Biosystems for Karyotyping and FISH
[hide]
v
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Cytogenetics: chromosomes
Basic
concepts
Karyotype
Ploidy
Genetic material/Genome
Chromatin
Euchromatin
Heterochromatin
Chromosome
Chromatid
Nucleosome
Nuclear organization
Types
Autosome/Sex chromosome (or allosome or heterosome)
Macrochromosome/Microchromosome
Circular chromosome/Linear chromosome
Extra chromosome (or accessory chromosome)
Supernumerary chromosome
A chromosome/B chromosome
Lampbrush chromosome
Polytene chromosome
Dinoflagellate chromosomes
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Centromere number
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Processes
and evolution
Mitosis
Meiosis
Structural alterations
Chromosomal inversion
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Numerical alterations
Aneuploidy
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Polyploidization
Structures
Telomere: Telomere-binding protein (TINF2)
Protamine
Histone
H1
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H2B
H3
H4
Centromere
A
B
C1
C2
E
F
H
I
J
K
M
N
O
P
Q
T
See also
Extrachromosomal DNA
Plasmid
List of organisms by chromosome count
List of chromosome lengths for various organisms
List of sequenced genomes
International System for Human Cytogenetic Nomenclature
Retrieved from "https://en.wikipedia.org/w/index.php?title=Karyotype&oldid=826784999"
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Cariotipo – Spanish
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Kariotipo – Basque
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Caryotype – French
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Cariotipo – Galician
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Kariotip – Croatian
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Cariotipo – Italian
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Karyotype – Dutch
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Karyotype – Norwegian
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Kariotyp – Polish
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Cariotip – Romanian
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Kariotip – Serbo-Croatian
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Karyotyp – Swedish
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