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Disc golf
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Disc golf
Highest governing body
Professional Disc Golf Association
Registered players
103,785 current[1]
Clubs
Yes
Characteristics
Contact
No
Team members
Single competitors, doubles
Mixed gender
Yes, but usually in separate leagues/divisions
Type
Outdoor
Equipment
Flying disc
Disc golf[2] (also called Frisbee Golf[3] or sometimes Frolf[4]) is a flying disc sport in which players throw a disc at a target, and is played using rules similar to golf.[5] It is often played on a course of 9 or 18 holes, but other formats are common. Players complete a hole by throwing a disc from a tee area toward a target, throwing again from the landing position of the disc until the target is reached. Players seek to complete a course in the lowest number of total throws.[5]
The game is played in about 40 countries and according to a 2017 study there are about 44,000 active members of the PDGA worldwide.[6]
Contents
[hide]
1 Origin and early history
1.1 Early pioneers of the sport
1.2 Creation of the modern game
2 Disc golf courses
2.1 Tees
2.2 Signs
2.3 Targets
3 Play of the game
4 Disc types
4.1 Putter
4.2 Mid-range
4.3 Driver
4.4 Stability
4.5 Plastics
5 Throwing styles
5.1 Backhand
5.2 Forehand
5.3 Alternative throws
6 Scoring
7 Tournaments
8 Popularity
8.1 Women in disc golf
9 Disc golf hall of fame
10 See also
11 External links
12 References
Origin and early history[edit]
Disc golf was first invented in the early 1900s. The first game was held in Bladworth, Saskatchewan, Canada in 1926.[7] Ronald Gibson and a group of his Bladworth Elementary School buddies played a game of throwing tin lids into 4 foot wide circles drawn into sandy patches on their school grounds.[8] They called the game Tin Lid Golf and played on a fairly regular basis. However, after they grew older and went their separate ways, the game came to an end.[9] It was not until the 1970s that disc golf would be reintroduced to Canadians at the Canadian Open Frisbee Championships in Toronto.[10][11]
Modern disc golf started in the early 1960s, but there is debate over who came up with the idea first. The consensus is that multiple groups of people played independently throughout the 1960s. Students at Rice University in Houston, Texas, for example, held tournaments with trees as targets as early as 1964, and in the early 1960s, players in Pendleton King Park in Augusta, Georgia would toss Frisbees into 50-gallon barrel trash cans designated as targets. In 1968 Frisbee Golf was also played in Alameda Park in Santa Barbara, Ca. by teenagers in the Anacapa and Sola street areas. Gazebos, water fountains, lamp posts, and trees were all part of the course. This took place for several years and an Alameda Park collectors edition disc still exists, though rare, as few were made. Clifford Towne from this group went on to hold a National Time Aloft record.
Early pioneers of the sport[edit]
Two early coordinators of the sport are George Sappenfield and Kevin Donnelly, who, through similar backgrounds and the help of Wham-O, were able to individually spread the sport in their California cities. Donnelly began playing a form of Frisbee golf in 1959 called Street Frisbee Golf. In 1961, while a recreation leader and then recreation supervisor for the City of Newport Beach, California, he formulated and then began organizing Frisbee golf tournaments at nine of the city's playgrounds he supervised. This culminated in 1965 with a fully documented, Wham-O sponsored, citywide Frisbee golf tournament. This highly publicized tournament included hula hoops as holes, with published rules, hole lengths, pars, and prizes; an event in which Walter Frederick Morrison, the Frisbee inventor, was in attendance.[12] In 1965, Sappenfield was a recreation counselor during a summer break from college during which, he set up an object course for his children to play on. When he finished college in 1968, Sappenfield became the Parks and Recreation supervisor for Conejo Recreation and Park District in Thousand Oaks, California. Sappenfield planned a disc golf tournament as part of a recreation project and contacted Wham-O Manufacturing to ask them for help with the event. Wham-O supplied Frisbees for throwing, and hula hoops for use as targets.[13] Before 1973 and the invention of the disc golf target called the disc pole hole, there were only a few disc golf object courses in the U.S. and Canada. Despite having never heard of the International Frisbee Association (IFA) that Ed Headrick and Wham-O had put together, or ever seeing a copy of the IFA Newsletter, Jim Palmeri, his brother, and a small group of people from Rochester, NY, had been playing disc golf as a competitive sport on a regular basis since August of 1970, including tournaments and weekly league play. By 1973, they had even promoted two City of Rochester Disc Frisbee Championship events which featured disc golf as the main event. In Canada, beginning in 1970, newly arrived Americans, Ken Westerfield and Jim Kenner played Frisbee golf daily on an 18 object hole course they designed at Queens Park in downtown Toronto. In California, the Berkeley Frisbee Group established a standardized 18 hole object course on the Berkeley campus in 1970. University of Michigan Nichols Arboretum, Ann Arbor had an object Frisbee golf course designed in the early 1970’s.[14][15]
Creation of the modern game[edit]
"Steady Ed" Headrick and Dave Dunipace are two inventors and players who greatly impacted how disc golf is played. Headrick introduced the first formal disc golf target with chains and a basket,[16] and Dave Dunipace invented the modern golf disc in 1983, with the revolutionary change of adding a beveled rim, giving the disc a greater distance and accuracy.[17] Dave was one of the founders of Innova, a well-known disc manufacturer.[18] In 1976, Headrick formed the DGA, then later the PDGA and the RDGA. Ted Smethers took over the PDGA in 1982 to be run independently and to officiate the standard rules of play for the sport.
"Steady Ed" Headrick began thinking about the sport during his time at Wham-O Toys where he designed and patented the modern day Frisbee.[19][20] Headrick designed and installed the first standardized target course in what was then known as Oak Grove Park in La Cañada Flintridge, California. (Today the park is known as Hahamongna Watershed Park). Ed worked for the San Gabriel, California-based Wham-O Corporation and is credited for pioneering the modern era of disc sports. Ed Founded "The International Frisbee Association (IFA)" and began establishing competitive standards for various sports using the Frisbee such as Distance, MTA, TRC, Accuracy, Freestyle, and Guts.
Headrick coined and trademarked the term "Disc Golf" when formalizing the sport and patented the Disc Pole Hole,[21] the first disc golf target to incorporate chains and a basket on a pole. He started designing the target because he was tired of arguing over what counted as a scoring disc with his friends. He founded the Professional Disc Golf Association (PDGA) and Recreational Disc Golf Association (RDGA) for competitive and family-oriented play, respectively, and worked on standardizing the rules and the equipment for the growing sport. Headrick abandoned his trademark on the term "Disc Golf", and turned over control and administration of the PDGA to the growing body of disc golf players in order to focus on his passion for building and inventing equipment for the sport.[22]
Disc golf courses[edit]
Disc golf courses usually have nine or 18 holes, and exceptions most often have holes in multiples of three.[23] Holes are designed to require a range of different throws to challenge players with different strengths or particular skills. According to the DGA, an average course hole should range around 200–240 feet (61–73 m) per hole.[24] Course designers use trees, bushes, elevation changes, water hazards, and distance variation, along with out-of-bounds zones and mandatory flight paths, to make each hole challenging and unique. Many courses include multiple tee positions or multiple target positions to cater to players of different ability levels.[23]
Disc golf courses are usually built in more natural and less manicured environments than golf and require much less maintenance. Professional course designers consider safety a critical factor in course design, and are careful to minimize the danger of being hit by a flying disc. Many courses are central organizing points for local disc golf clubs, and some include shops selling disc golf equipment. More than 80% of the courses listed on dgcoursereview.com are listed as public and free to play.[25]
Tees[edit]
A disc golf tee position is usually a pad of concrete, asphalt, rubber, gravel, or artificial turf. The PDGA recommends that they be no smaller than 1.2 meters wide by 3 meters long.[23] Many courses have only the front of the tee position marked with a board of timber in the ground. Some courses have no tees at all.[citation needed]
Signs[edit]
Established courses have tee signs near each tee position, showing a simple map of the hole, distance to the hole, par, out-of-bounds areas, mandatory paths, and a unique name for the hole. They are often supplemented with a larger sign near the course entrance which has a course map and other information.[23]
Targets[edit]
A red disc sailing towards a "Tonal Pole" style target at the disc golf course on Pender Island.
Although early courses were played using trees, fence posts, or park equipment as the target, modern courses typically use tone targets or standard disc golf baskets. Tone targets are designed to make a distinctive sound when hit with a disc. Baskets are now much more common, which are constructed with a central pole holding a basket under an assembly of hanging chains. A disc which hits the chains is often, but not always, deflected into the basket. Discs must land inside the basket or on the chains to count the hole as completed. There are many different brands of baskets made by numerous manufacturers.
Play of the game[edit]
The sport of disc golf is set up similar to a game of golf. A "round" is played on a disc golf course consisting of a number of "holes", usually 9 or 18. Each hole includes a tee position for starting play and a disc golf target some distance away, often with obstacles such as trees, hills or bodies of water in between.[26] Players begin by throwing a disc from the tee, and navigate the hole by picking up the disc where it lands and throwing again until they reach the target. The object of the game is to get through the course with the lowest number of total throws.[27] Play is usually in groups of five or fewer, with each player taking turn at the tee box, then progressing with the player furthest from the hole throwing first, while the other players stand aside.
Each course is unique, so each course requires a different combination of throws to complete, with the best players aiming to shape the flight of the disc to account for distance, terrain, obstacles and weather. In order to facilitate making different shots, players carry a variety of discs with different flight characteristics, choosing an appropriate disc for each throw. Some players also carry a mini marker disc, used to accurately mark the throwing position before each throw. Use of mini marker discs is particularly prevalent in formal competitive play.
Many courses include out-of-bounds areas, commonly called "OB zones" or just "OB". If the disc lands in these areas, the player is usually required to add a penalty throw onto his or her score and continue play from near where the disc entered the out-of-bounds zone. Some courses include out-of-bounds areas with special rules requiring the player the resume play from a specified area called a drop zone, or requiring the player to restart the hole from the tee. Some courses also include Mandatories (also called "Mandos") which require the path of the disc to be above, below or to one side of a specific line indicated by a sign.
By tradition, players throw from the tee box in the order of their score on the previous hole, with the lowest scorer throwing first. Most players also follow a loose code of courtesy while playing, which includes norms such as standing out of the sight line of the throwing player and avoiding making distracting noises. Because a thrown disc could injure someone, the Professional Disc Golf Association recommends that players "Never throw into a blind area or when spectators, pedestrians or facility users are within range."[28]
Formal competitive play is governed by the PDGA Official Rules of Disc Golf and the PDGA Competition Manual for Disc Golf events.
A disc resting in the basket
Disc types[edit]
The golf discs used today are much smaller and heavier than traditional flying discs, typically 8–9 inches (20–23 cm) in diameter and weighing between 120 and 180 grams. The PDGA prohibits discs heavier than 200 grams. Discs used for disc golf are designed and shaped for control, speed, and accuracy, while general-purpose flying discs, such as those used for playing guts or ultimate, have a more traditional shape, similar to a catch disc. There is a wide variety of discs used in disc golf and they are generally divided into three categories: putters, all-purpose mid-range discs, and drivers.
Putter[edit]
Putters are similar to the discs used in simple games of catch, such as the Wham-o brand Frisbee. They are designed to fly straight, predictably, and very slowly compared to mid-range discs and drivers. They are typically used for tight, controlled shots that are close to the basket, although some players use them for short drives where trees or other obstacles come into play. Usually a pro carries 1–7 putters depending on their flight characteristics.
Mid-range[edit]
Mid-range discs have slightly sharper edges that enable them to cut through the air better. These discs are usually faster, more stable, and have a longer range than a putter. Some players will use mid-ranges as drivers, and there are tournaments that require players to use only mid-range discs. They are good all-around discs and are suitable for a first time player.
Driver[edit]
Drivers are usually recognized by their sharp, beveled edge and have most of their mass concentrated on the outer rim of the disc rather than distributed equally throughout. They are designed to travel farther distances at greater speeds and are mostly used for tee-off and other long distance throws.[29] Drivers are often divided into different categories. For example, Innova Discs divides their discs into Distance Drivers and Fairway Drivers, with a fairway driver being somewhere between a distance driver and a mid-range disc. Discraft divides their drivers into 3 categories: Long Drivers, Extra Long Drivers, and Maximum Distance Drivers. Because the physics of a disc require "snap" or "flick", which means putting spin on the disc, new players generally find that throwing a distance driver accurately can be somewhat difficult and will require experience with golf disc response. This is why it is better for players to begin with fairway drivers, long drivers, or even mid-ranges, and incorporate maximum distance drivers as their strength and disc control increases. Most players that are starting off will be most likely throwing lighter discs. Another type of driver, used less frequently, is a roller. As the name indicates, it has an edge designed to roll rather than fly.
Stability[edit]
Stability is the measurement of a disc's tendency to bank laterally during its flight. A disc that is over-stable will tend to track left (for a right handed, backhand throw), whereas a disc that is under-stable will tend to track right (also for a right handed, backhand throw). The stability rating of the discs differs depending on the manufacturer of the disc. Innova Discs rate stability as "turn" and "fade". "Turn" references how the disc will fly at high speed during the beginning and middle of its flight, and is rated on a scale of +1[30] to −5, where +1[30] is the most overstable and −5 is the most understable. "Fade" references how the disc will fly at lower speeds towards the end of its flight, and is rated on a scale of 0 to 5, where 0 has the least fade, and 5 has the most fade. For example, a disc with a turn of −5 and fade of +1 will fly to the right for (right handed, backhand throw) the majority of its flight then curl back minimally left at the end. A disc with a turn of −1 and a fade of +3 will turn slightly right during the middle of its flight and turn hard left as it slows down. These ratings can be found on the discs themselves or from the manufacturer's web site. Discraft prints the stability rating on all discs and also provides this information on their web site. The stability ranges from 3 to −2 for Discraft discs; however Discraft's ratings are more of a combination of turn and fade with the predominance being fade.
Spin (rotation) has little influence on lift and drag forces but impacts a disc's stability during flight. Imagine a spinning top. A gentle nudge will knock it off its axis of rotation for a second, but it will not topple over because spin adds gyroscopic stability. In the same way, a flying disc resists rolling (flipping over) because spin adds gyroscopic stability. A flying disc will maintain its spin rate even as it loses velocity. Toward the end of a disc's flight, when the spin and velocity lines cross, a flying disc will predictably begin to fade. The degree to which a disc will fade depends on its pitch angle and design.[31]
Plastics[edit]
There are a variety of different discs, each with a specific plastic made with them. Plastics such as DX, J-Pro, Pro-D, X-Line and R-Pro from Innova discs and Discraft are some of the less durable plastics, but good for beginners due to their lower prices, compared to the higher end plastics. Plastics such as Champion, Titanium, FLX, GStar, and Star plastics, which are the best offered from the same companies, offering the best quality, durability and flight compared to the other types available.
Throwing styles[edit]
Main article: Flying disc techniques
While there are many different grips and styles to throwing the disc, there are two basic throwing techniques: backhand and forehand (or sidearm). These techniques vary in effectiveness under different circumstances. Their understanding and mastery can greatly improve a player's game, and offer diverse options in maneuvering the disc to the basket with greater efficacy. Many players use what is referred to as a run-up during their drive. This is practiced to build more forward disc momentum and distance. Throwing styles vary from player to player, and there is no standard throwing style.
All discs when thrown will naturally fall to a certain direction, this direction is termed Hyzer, the natural fall of the disc, or Anhyzer, making the disc fall against its natural flight pattern. For a right-handed backhand throw (RHBH), the disc will naturally fall to the left. For a right-handed forehand throw (RHFH), the disc will naturally fall to the right. For a left-handed, backhand throw (LHBH), the disc will naturally fall to the right. For a left-handed, forehand throw (LHFH), the disc will naturally fall to the left.
Backhand[edit]
To perform this throw, the disc is rapidly drawn from across the front of the body, and released towards a forward aimpoint. Due to the potential snap available with this technique, one can expect greater distance and accuracy than with a forehand throw. It is important to initiate momentum from the feet and allow it to travel up the body, hips and shoulders, culminating in the transfer of energy to the disc.
Forehand[edit]
The forehand (sidearm) throw is performed by drawing the disc from behind and partially across the front of the body: similar to a sidearm throw in baseball. The term sidearm actually predates the term forehand, which is seemingly in use today as a simpler means to communicate the technique, equating to a tennis forehand.[32]
Alternative throws[edit]
The following examples of throws may be used to better deliver a disc where the former common two throws would be impeded by obstacles such as bushes, trees, boulders, or artificial structures.
Common alternative styles
The Hatchet (or Tomahawk). Gripped similarly to the sidearm toss but thrown with an overhand motion; the disc orientation nearly perpendicular to the ground over much of the flight.
The Thumber (or U.D.). Thrown in an overhand manner but with thumb held on the disc's underside.
The Roller. Thrown either backhand or forehand, the disc will predominately be in contact with the ground. The disc remains in motion while travelling on its edge at a slight angle, and can travel exceedingly far in ideal situations. Once perfected, the roller is an invaluably versatile tool in the golfer's arsenal.
The Turbo-Putt Thrown with a putter when the player holds the disc upright, supported in the middle by the thumb, with the finger tips outside of the edge, somewhat like a waiter holding a platter. The player stands with the leg opposite from the throwing arm forward, reaches back, and then extends their arm towards the basket, throwing the disc in a motion similar to that of throwing a dart. Ideally the thrower does not rotate his wrist; the act of following through will give the disc its spin.[33] The Turbo-Putt is a throw known for its accuracy, but it has extremely limited range.
Other alternative styles
The Baseball or Grenade. Thrown as in the backhand, but with the disc upside-down. This shot is used often to get up and down on a short shot where there is danger of a shot rolling away or going out of bounds if thrown too far. Primarily used on downhill shots but can be used to go up and over. Also due to the quick turn and backspin of this shot, it is sometimes used to get out of the woods.
The Overhand wristflip (or chicken-wing [ambiguous origin][dubious – discuss] ). This is a very difficult and stylized throw with which accomplished free-stylers and classic ultimate players are familiar; it is less used in disc golf. It is thrown in the same manner the "baseball" but drawn on the sidearm side of the body, and by inverting the arm and disc. Using the thumb as the power finger, the disc is drawn from the thigh area rearwards and up from behind the body to over the shoulder, releasing toward a forward aimpoint. The disc flies in a conventional flight pattern. To the untrained eye, this appears to be an ungainly throw. It is, however, elegant and accurate. The term "overhand wristflip" has been in use since at least circa 1970.
Scoring[edit]
Stroke Play is the most common scoring method used in the sport but there are many other forms. These include match play, skins, speed golf and captain's choice, which in disc golf is referred to as "doubles" (not to be confused with partner or team play).
Regardless of which form of play the participants choose, the main objectives of disc golf are conceptually the same as traditional golf in the sense that players follow the same scorekeeping technique.
Scoreboard:
Condor – Where a player is four throws under par, or "-4".
Albatross (or double-eagle) – Where a player is three throws under par, or "-3".
Eagle (or double-birdie) – Where a player is two throws under par, or "-2".
Birdie – Where a player is one throw under par, or "-1".
Par – Where a player has thrown par, or "0".
Bogey – Where a player is one throw over par, or "+1".
Double Bogey – Where a player is two throws over par, or "+2".
Triple Bogey – Where a player is three throws over par, or "+3".
Doubles play is a unique style of play that many local courses offer on a weekly basis. In this format, teams of two golfers are determined. Sometimes this is done by random draw, and other times it is a pro-am format. On the course, it is a "best-disc" scramble, meaning both players throw their tee shot; and then decide which lie they would like to play. Both players then play from the same lie, again choosing which lie is preferable. The World Amateur Doubles Format includes best shot, alternate shot, best score (players play singles and take the best result from the hole) and worst shot (both players must sink the putt).
Tournaments[edit]
Ken Climo teeing off at hole 5 of the 2008 USDGC
Tournaments are held nationwide and year long in the United States. Sanctioned Tournament play is communicated through the Professional Disc Golf Association Membership. The PDGA provides international, professional, and amateur disc golf tournaments as well as communicates event results, opinions and other information beneficial to the sport via electronic and printed media. In 1982 the PDGA hosted the first World Championship Tournament. Since then, the World Championships have been held in 17 different American states, as well as Toronto, Ontario.[34]
Disc golf tournaments are popular around the world. As with traditional golf, there are many championship tournaments. One of the largest is the United States Disc Golf Championship.
Every year, the largest teams tournament in the world is held in Austin, Texas, by John Houck.[35]
To prove the year-round sustainability of the sport, annual winter tournaments known as Ice Bowls are held at courses around the world. Using the motto "No Wimps, No Whiners", Ice Bowls collectively are designed to create sport awareness, and are considered charity events that typically benefit a food bank local to a given tournament location. The official Web site reports that the 2010 Ice Bowls raised over $250,000 and donated over 67,000 pounds of food in the 222 tournaments for the year. [2]
Popularity[edit]
Disc golf is a rapidly growing sport worldwide,[36] and is the 4th fastest growing sport in United States, behind MMA, Roller Derby, and Parkour.[37] DGCourseReview.com, which tracks courses worldwide along with opening dates, shows a rapid increase in installed permanent courses with an average of more than 400 new courses added each year between 2007 and 2017. The site lists 6800 courses worldwide (in May 2017).[38]
Although most players play on a casual, amateur level, the professional disc golf scene is also growing rapidly, with the top professionals playing full-time and earning their livings through tournament winnings and sponsorship from equipment manufacturers. Online viewership of major tournaments and events has increased rapidly, with coverage of several tournaments in 2016 achieving more than 75,000 views on YouTube.[39][40] and a clip focused on a single throw by professional Philo Braithwaite achieving more than 1.1 million views.
Women in disc golf[edit]
While there are more male than female players, the Women's Disc Golf Association exists to encourage female players and arrange women's tournaments. A PDGA survey states that out of its 35,662 active members in 2016, 7.6% are female, or about 2,728.[6]
Several companies have started programs and websites to help attract women to the sport. The PDGA Women's Committee is "Dedicated to Attract, Encourage, and Retain Female Participation in Organized Disc Golf Events". The PDGA Women's Committee set historical records on 12 May 2012 by running the Inaugural Women's Global Event that attracted 636 female players in 24 states and 4 countries. The Women's Global Event was expected to take place every two years from 2014, with hopes of increasing the number of participants.
There are also Disc golf companies such as Disc-Diva, that have started up with a primary, though not exclusive, focus on women in the sport, promoting accessories geared towards women and using catch phrases like "you wish you threw like a girl".[41] Sassy Pants is another group that focuses on getting more involvement from women in the sport, advocating for sponsorship of women to enter tournaments.[42]
Women's disc golf teams are involved in the National Collegiate Disc Golf Championship, and the Mississippi State Women's Team were the inaugural champions.
Disc golf hall of fame[edit]
Inductees:[43]
1993 Vanessa Chambers | Dave Dunipace | Ed Headrick | Tom Monroe | Jim Palmeri | Dan Roddick | Ted Smethers
1994 Harold Duvall | Nobuya Kobayashi | Darrell Lynn | Dan Mangone | Doug Newland | Snapper Pierson | Lavone Wolfe
1995 Ken Climo | John David | David Greenwell | Johnny Roberts | Dr. Rick Voakes
1996 Mike Conger | Patti Kunkle | Rick Rothstein
1997 Steve Slasor | Elaine King | Jim Kenner
1998 Gregg Hosfeld | John Houck | Carlton Howard
1999 Sam Ferrans | Steve Wisecup | Tim Selinske
2000 Tom Schot | Royce Racinowski
2001 Stan McDaniel | Johnny Sias
2002 Alan Beaver | Gary Lewis
2003 Mark Horn | Brian Hoeniger | Dr. Stancil Johnson,
2004 Derek Robins | Geoff Lissaman | Johnny Lissaman | Marty Hapner
2005 Mats Bengtsson | Sylvia Voakes
2006 Chuck Kennedy | Kozo Shimbo
2007 Fred Salaz | Michael Travers
2008 Dan Ginnelly | Juliana Korver
2009 Crazy John Brooks | Lynne Warren | Michael Sullivan
2010 Charlie Callahan | Tomas Ekstrom | Brian Cummings
2011 Don Hoffman | Joe Feidt | Brent Hambrick
2012 Tim Willis | Jeff Homburg | Bob Gentil (New Zealand)
2013 Barry Schultz | Becky Zallek | Jim Challas | Ken Westerfield
2014 Don Wilchek | Jim Oates
2015 Gail McColl | Anni Kremi | J Gary Dropcho
2016 Joseph Mela | Ace Mason | Tita Ugalde
See also[edit]
Flying disc games
Flying disc techniques
Frisbee
Golf
External links[edit]
Wikimedia Commons has media related to Disc golf.
Wikibooks has a book on the topic of: Disc Golf
Professional Disc Golf Association (PDGA) – official rules, course directory and list of members
History of Early Frisbee Sports
History of Disc Golf, Ultimate and Frisbee Pioneers
References[edit]
Jump up ^ "PDGA Player Search". PDGA. Retrieved 2016-04-06.
Jump up ^ "What is Disc Golf - DGA | Disc Golf Association". DGA | Disc Golf Association. Retrieved 2017-05-06.
Jump up ^ "Definition of disc golf". Retrieved 2017-12-20.
Jump up ^ "Is The Hand-Wringing Over 'Frolf' Worth The Worry?". Ultiworld Disc Golf. Retrieved 20 December 2017.
^ Jump up to: a b "On Your Own; Connecting a Fairway". The New York Times. 1990-07-02. ISSN 0362-4331. Retrieved 2017-05-10.
^ Jump up to: a b "2016 PDGA Membership Growth" (PDF). March 15, 2017.
Jump up ^ "History of Disc Golf – Deaf Disc Golf Association". www.ddga.org. Retrieved 2017-06-02.
Jump up ^ Pg. 72" A Chain of Events – The Origin and Evolution of Disc Golf"
Jump up ^ "Disc Golf". Disc Golf History. Retrieved 28 June 2013.
Jump up ^ "Canadian Open Frisbee Championships". Frisbee Pioneer Ken Westerfield. Retrieved 24 December 2014.
Jump up ^ "History of Frisbee and Flying Disc freestyle". Development of Frisbee in Canada. Retrieved October 16, 2017.
Jump up ^ Altmyer, Don (2011). "Kevin Donnelly: Disc Golf". Disc Golfer (Fall 2011).
Jump up ^ Gregoire, Matt (2008-09-23). "Brief History of Disc Golf and the PDGA". Professional Disc Golf Association. Retrieved 2017-05-11.
Jump up ^ "Greater Rochester Disc Golf Club". Retrieved December 25, 2017.
Jump up ^ "History of Ultimate Frisbee and Disc Sports". Retrieved December 25, 2017.
Jump up ^ "About Ed Headrick - Father of Disc Golf - DGA | Disc Golf Association". DGA | Disc Golf Association. Retrieved 2017-05-11.
Jump up ^ Gregoire, Matt (2015-01-30). "Disc Golf Hall Of Fame". Professional Disc Golf Association. Retrieved 2017-05-11.
Jump up ^ "Innova - The Choice Of Champions - #1 in Disc Golf - Innova Disc Golf". innovadiscs.com. Retrieved 2017-05-11.
Jump up ^ "Ed Headrick, Designer of the Modern Frisbee, Dies at 78". New York Times. Retrieved 2002-06-14.
Jump up ^ "The History of Disc Golf". Discgolf.com. Retrieved 2011-12-27.
Jump up ^ US 4039189
Jump up ^ "Archive: 1978 Disc Golf Association Disc Golf Promotional Development Guide. PDF 11 pages" (PDF). Discgolf.com. Retrieved 2013-09-29.
^ Jump up to: a b c d "PDGA Disc Golf Course Design Recommendations" (PDF). PDGA.com.
Jump up ^ "PDGA Disc Golf Course Design Standards - DGA | Disc Golf Association". DGA | Disc Golf Association. Retrieved 2017-05-11.
Jump up ^ "Disc Golf Course Review: View and review over 6000 Disc Golf Courses!". www.dgcoursereview.com. Retrieved 2017-05-09.
Jump up ^ Slocum, Bill (2000-06-11). "A Good Golf Game? It's All in the Wrist". The New York Times. ISSN 0362-4331. Retrieved 2017-05-10.
Jump up ^ pdga (2008-10-09). "A Guide to Disc Golf from the PDGA". Professional Disc Golf Association. Retrieved 2017-05-09.
Jump up ^ steveganz (2016-01-20). "The Disc Golfer's Code". Professional Disc Golf Association. Retrieved 2017-05-09.
Jump up ^ Eastham, Susan L. "DISC GOLF: Teaching A Lifetime Activity." Strategies (08924562) 28.6 (2015): 3–8. SPORTDiscus with Full Text. Web. 1 Aug. 2016.
^ Jump up to: a b http://www.innovadiscs.com/home/disc-golf-faq/flight-ratings-system.html
Jump up ^ "The Definitive Guide to Disc Golf". www.triumphbooks.com. Retrieved 2016-04-21.
Jump up ^ "Throwing sidearm". Disc golf review. Retrieved 6 January 2015.
Jump up ^ https://www.youtube.com/watch?v=wzA8y0rwGuo
Jump up ^ http://www.bestdiscgolfdiscs.com/infographic-pdga-world-championship/
Jump up ^ "Texas Disc Golf Team Championships". Circular Productions. Retrieved 2011-12-27.
Jump up ^ "2007-2014 International Demographics" (PDF).
Jump up ^ Golf, Parked: The Sociology of Disc (2017-05-01). "Disc golf is trending in U.S. newspapers: Supporting evidence for disc golf's "fast-growing sport" claim". Parked. Retrieved 2017-05-11.
Jump up ^ "Disc Golf Courses in the World - Disc Golf Course Review". www.dgcoursereview.com. Retrieved 2017-05-11.
Jump up ^ "thediscgolfguy". YouTube. Retrieved 2017-05-11.
Jump up ^ "Central Coast Disc Golf". YouTube. Retrieved 2017-05-11.
Jump up ^ "Disc Diva Socks". Discgolfdiva.startlogic.com. Retrieved 2013-09-29.
Jump up ^ [1] Archived 27 July 2014 at the Wayback Machine.
Jump up ^ "Disc Golf Hall of Fame". Professional Disc Golf Association. Retrieved 15 April 2016.
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Eukaryotic transcription
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This article's lead section does not adequately summarize key points of its contents. Please consider expanding the lead to provide an accessible overview of all important aspects of the article. Please discuss this issue on the article's talk page. (November 2016)
Eukaryotic transcription is the elaborate process that eukaryotic cells use to copy genetic information stored in DNA into units of RNA replica. Gene transcription occurs in both eukaryotic and prokaryotic cells. Unlike prokaryotic RNA polymerase that initiates the transcription of all different types of RNA, RNA polymerase in eukaryotes (including humans) comes in three variations, each encoding a different type of gene. A eukaryotic cell has a nucleus that separates the processes of transcription and translation. Eukaryotic transcription occurs within the nucleus where DNA is packaged into nucleosomes and higher order chromatin structures. The complexity of the eukaryotic genome necessitates a great variety and complexity of gene expression control.
Contents
[hide]
1 Overview
2 RNA polymerase
3 Initiation
3.1 Eukaryotic promoters and general transcription factors
3.2 Assembly of preinitiation complex
3.3 Promoter melting and open complex formation
3.4 Abortive initiation
3.5 Promoter escape
4 Elongation
4.1 Elongation factors
4.2 Transcription fidelity
4.3 Pausing, poising, and backtracking
4.4 RNA processing
5 Termination
5.1 Factor-dependent
5.2 Factor-independent
6 Eukaryotic transcriptional control
6.1 Global control and epigenetic regulation
6.2 Gene-specific activation
6.3 Gene-specific repression
6.4 Elongation and termination control
7 Transcription-coupled DNA repair
8 Comparisons between prokaryotic and eukaryotic transcription
9 Regulation of transcription in cancer
10 References
Overview[edit]
Transcription is the process of copying genetic information stored in a DNA strand into a transportable complementary strand of RNA.[1] Eukaryotic transcription takes place in the nucleus of the cell and proceeds in three sequential stages: initiation, elongation, and termination.[1] The transcriptional machinery that catalyzes this complex reaction has at its core three multi-subunit RNA polymerases. RNA polymerase I is responsible for transcribing RNA that codes for genes that become structural components of the ribosome. [1]
Protein coding genes are transcribed into messenger RNAs (mRNAs) that carry the information from DNA to the site of protein synthesis.[1] Although mRNAs possess great diversity, they are not the most abundant RNA species made in the cell. The so-called non-coding RNAs account for the large majority of the transcriptional output of a cell.[2] These non-coding RNAs perform a variety of important cellular functions.[2]
RNA polymerase[edit]
Eukaryotes have three nuclear RNA polymerases, each with distinct roles and properties[3][4]
Name Location Product
RNA Polymerase I (Pol I, Pol A) nucleolus larger ribosomal RNA (rRNA) (28S, 18S, 5.8S)
RNA Polymerase II (Pol II, Pol B) nucleus messenger RNA (mRNA), most small nuclear RNAs (snRNAs), small interfering RNA (siRNAs) and microRNA (miRNA).
RNA Polymerase III (Pol III, Pol C) nucleus (and possibly the nucleolus-nucleoplasm interface) transfer RNA (tRNA), other small RNAs (including the small 5S ribosomal RNA (5s rRNA), snRNA U6, signal recognition particle RNA (SRP RNA) and other stable short RNAs
RNA polymerase I (Pol I) catalyses the transcription of all rRNA genes except 5S.[3][4] These rRNA genes are organised into a single transcriptional unit and are transcribed into a continuous transcript. This precursor is then processed into three rRNAs: 18S, 5.8S, and 28S. The transcription of rRNA genes takes place in a specialised structure of the nucleus called the nucleolus,[5] where the transcribed rRNAs are combined with proteins to form ribosomes.[6]
RNA polymerase II (Pol II) is responsible for the transcription of all mRNAs, some snRNAs, siRNAs, and all miRNAs.[3][4] Many Pol II transcripts exist transiently as single strand precursor RNAs (pre-RNAs) that are further processed to generate mature RNAs.[1] For example, precursor mRNAs (pre-mRNAs)are extensively processed before exiting into the cytoplasm through the nuclear pore for protein translation.
RNA polymerase III (Pol III) transcribes small non-coding RNAs, including tRNAs, 5S rRNA, U6 snRNA, SRP RNA, and other stable short RNAs such as ribonuclease P RNA.[7]
Structure of eukaryotic RNA polymerase II (light blue) in complex with α-amanitin (red), a strong poison found in death cap mushrooms that targets this vital enzyme
RNA Polymerases I, II, and III contain 14, 12, and 17 subunits, respectively.[8] All three eukaryotic polymerases have five core subunits that exhibit homology with the β, β’, αI, αII, and ω subunits of E. coli RNA polymerase. An identical ω-like subunit (RBP6) is used by all three eukaryotic polymerases, while the same α-like subunits are used by Pol I and III. The three eukaryotic polymerases share four other common subunits among themselves. The remaining subunits are unique to each RNA polymerase. The additional subunits found in Pol I and Pol III relative to Pol II, are homologous to Pol II transcription factors.[8]
Crystal structures of RNA polymerases I[9] and II [10] provide an opportunity to understand the interactions among the subunits and the molecular mechanism of eukaryotic transcription in atomic detail.
The carboxyl terminal domain (CTD) of RPB1, the largest subunit of RNA polymerase II, plays an important role in bringing together the machinery necessary for the synthesis and processing of Pol II transcripts.[11] Long and structurally disordered, the CTD contains multiple repeats of heptapeptide sequence YSPTSPS that are subject to phosphorylation and other posttranslational modifications during the transcription cycle. These modifications and their regulation constitute the operational code for the CTD to control transcription initiation, elongation and termination and to couple transcription and RNA processing.[11]
Initiation[edit]
The initiation of gene transcription in eukaryotes occurs in specific steps.[1] First, an RNA polymerase along with general transcription factors binds to the promoter region of the gene to form a closed complex called the preinitiation complex. The subsequent transition of the complex from the closed state to the open state results in the melting or separation of the two DNA strands and the positioning of the template strand to the active site of the RNA polymerase. Without the need of a primer, RNA polymerase can initiate the synthesis of a new RNA chain using the template DNA strand to guide ribonucleotide selection and polymerization chemistry.[1] However, many of the initiated syntheses are aborted before the transcripts reach a significant length (~10 nucleotides). During these abortive cycles, the polymerase keeps making and releasing short transcripts until it is able to produce a transcript that surpasses ten nucleotides in length. Once this threshold is attained, RNA polymerase passes the promoter and transcription proceeds to the elongation phase.[1]
Here is a diagram of the attachment of RNA polymerase II to the de-helicized DNA. Credit: Forluvoft.
Eukaryotic promoters and general transcription factors[edit]
Pol II-transcribed genes contain a region in the immediate vicinity of the transcription start site (TSS) that binds and positions the preinitiation complex. This region is called the core promoter because of its essential role in transcription initiation.[12][13] Different classes of sequence elements are found in the promoters. For example, the TATA box is the highly conserved DNA recognition sequence for the TATA box binding protein, TBP, whose binding initiates transcription complex assembly at many genes.
Eukaryotic genes also contain regulatory sequences beyond the core promoter. These cis-acting control elements bind transcriptional activators or repressors to increase or decrease transcription from the core promoter. Well-characterized regulatory elements include enhancers, silencers, and insulators. These regulatory sequences can be spread over a large genomic distance, sometimes located hundreds of kilobases from the core promoters.[1]
General transcription factors are a group of proteins involved in transcription initiation and regulation.[1] These factors typically have DNA-binding domains that bind specific sequence elements of the core promoter and help recruit RNA polymerase to the transcriptional start site. General transcription factors for RNA polymerase II include TFIID, TFIIA, TFIIB, TFIIF, TFIIE, and TFIIH.[1][14][15]
Assembly of preinitiation complex[edit]
To prepare for transcription, a complete set of general transcription factors and RNA polymerase need to be assembled at the core promoter to form the ~2 million dalton preinitiation complex.[16] For example, for promoters that contain a TATA box near the TSS, the recognition of TATA box by the TBP subunit of TFIID initiates the assembly of a transcription complex. The next proteins to enter are TFIIA and TFIIB, which stabilize the DNA-TFIID complex and recruit Pol II in association with TFIIF and additional transcription factors. TFIIF serves as the bridge between the TATA-bound TBP and polymerase. One of the last transcription factors to be recruited to the preinitiation complex is TFIIH, which plays an important role in promoter melting and escape.[17]
The diagram describes the eukaryotic preinitiation complex which includes the general transcription factors and RNA Polymerase II. Credit: ArneLH.
Promoter melting and open complex formation[edit]
For pol II-transcribed genes, and unlike bacterial RNA polymerase, promoter melting requires hydrolysis of ATP and is mediated by TFIIH.[17] TFIIH is a ten-subunit protein, including both ATPase and protein kinase activities.[18] While the upstream promoter DNA is held in a fixed position by TFIID, TFIIH pulls downstream double-stranded DNA into the cleft of the polymerase, driving the separation of DNA strands and the transition of the preinitiation complex from the closed to open state. TFIIB aids in open complex formation by binding the melted DNA and stabilizing the transcription bubble.
Abortive initiation[edit]
Main article: Abortive initiation
Once the initiation complex is open, the first ribonucleotide is brought into the active site to initiate the polymerization reaction in the absence of a primer.[1] This generates a nascent RNA chain that forms a hetero-duplex with the template DNA strand. However, before entering the elongation phase, polymerase may terminate prematurely and release a short, truncated transcript. This process is called abortive initiation.[19] Many cycles of abortive initiation may occur before the transcript grows to sufficient length to promote polymerase escape from the promoter. Throughout abortive initiation cycles, RNA polymerase remains bound to the promoter and pulls downstream DNA into its catalytic cleft in a scrunching-kind of motion.[19]
Promoter escape[edit]
When a transcript attains the threshold length of ten nucleotides, it enters the RNA exit channel.[1] The polymerase breaks its interactions with the promoter elements and any regulatory proteins associated with the initiation complex that it no longer needs.[20] Promoter escape in eukaryotes requires ATP hydrolysis and, in the case of Pol II-phosphorylation of the CTD. Meanwhile, the transcription bubble collapses down to 12-14 nucleotides, providing kinetic energy required for the escape.[1]
Elongation[edit]
After escaping the promoter and shedding most of the transcription factors for initiation, the polymerase acquires new factors for the next phase of transcription: elongation.[21][22] Transcription elongation is a processive process. Double stranded DNA that enters from the front of the enzyme is unzipped to avail the template strand for RNA synthesis. For every DNA base pair separated by the advancing polymerase, one hybrid RNA:DNA base pair is immediately formed. DNA strands and nascent RNA chain exit from separate channels; the two DNA strands reunite at the trailing end of the transcription bubble while the single strand RNA emerges alone.
Elongation factors[edit]
Among the proteins recruited to polymerase are elongation factors, thus called because they stimulate transcription elongation.[23] There are different classes of elongation factors. Some factors can increase the overall rate of transcribing, some can help the polymerase through transient pausing sites, and some can assist the polymerase to transcribe through chromatin.[24] One of the elongation factors, P-TEFb, is particularly important.[25] P-TEFb phosphorylates the second residue (Ser-2) of the CTD repeats (YSPTSPS) of the bound Pol II. P-TEFb also phosphorylates and activates SPT5 and TAT-SF1. SPT5 is a universal transcription factor that helps recruit 5'-capping enzyme to Pol II with a CTD phosphorylated at Ser-5. TAF-SF1 recruits components of the RNA splicing machinery to the Ser-2 phosphorylated CTD. P-TEFb also helps suppress transient pausing of polymerase when it encounters certain sequences immediately following initiation.[25]
Transcription fidelity[edit]
Transcription fidelity is achieved through multiple mechanisms. RNA polymerases select correct nucleoside triphosphate (NTP) substrate to prevent transcription errors. Only the NTP which correctly base pairs with the coding base in the DNA is admitted to the active center.[26][27] RNA polymerase performs two known proof reading functions to detect and remove misincorporated nucleotides: pyrophosphorylytic editing and hydrolytic editing.[1] The former removes the incorrectly inserted ribonucleotide by a simple reversal of the polymerization reaction, while the latter involves backtracking of the polymerase and cleaving of a segment of error-containing RNA product. Elongation factor TFIIS stimulates an inherent ribonuclease activity in the polymerase, allowing the removal of misincorporated bases through limited local RNA degradation.[28] Note that all reactions (phosphodiester bond synthesis, pyrophosphorolysis, phosphodiester bond hydrolysis) are performed by RNA polymerase by using a single active center.[29]
Pausing, poising, and backtracking[edit]
Transcription elongation is not a smooth ride along the DNA railway. For proofreading, the polymerase is made to back-up, erase some of the RNA it has already made and have another go at transcription.[1] In general, RNA polymerase does not transcribe through a gene at a constant pace. Rather it pauses periodically at certain sequences, sometimes for long periods of time before resuming transcription.[30] In extreme cases, for example, when the polymerase encounters a damaged nucleotide, it comes to a complete halt. More often, an elongating polymerase is stalled near the promoter.[30] Promoter-proximal pausing during early elongation is a commonly used mechanism for regulating genes poised to be expressed rapidly or in a coordinated fashion. Pausing is mediated by a complex called NELF (negative elongation factor) in collaboration with DSIF (DRB-sensitivity-inducing factor containing SPT4/SPT5).[31] The blockage is released once the polymerase receives an activation signal, such as the phosphorylation of Ser-2 of CTD tail by P-TEFb. Other elongation factors such as ELL and TFIIS stimulate the rate of elongation by limiting the length of time that polymerase pauses.[1]
RNA processing[edit]
Elongating polymerase is associated with a set of protein factors required for various types of RNA processing.[1] mRNA is capped as soon as it emerges from the RNA-exit channel of the polymerase. After capping, dephosphorylation of Ser-5 within the CTD repeats may be responsible for dissociation of the capping machinery. Further phosphorylation of Ser-2 causes recruitment of the RNA splicing machinery that catalyzes the removal of non-coding introns to generate mature mRNA.[1] Alternative splicing expands the protein complements in eukaryotes. Just as with 5’-capping and splicing, the CTD tail is involved in recruiting enzymes responsible for 3’-polyadenylation, the final RNA processing event that is coupled with the termination of transcription.[1]
Termination[edit]
The last stage of transcription is termination, which leads to the dissociation of the complete transcript and the release of RNA polymerase from the template DNA.The process differs for each of the three RNA polymerases.[32] The mechanism of termination is the least understood of the three transcription stages.
Factor-dependent[edit]
The termination of transcription of pre-rRNA genes by polymerase Pol I is performed by a system that needs a specific transcription termination factor.[3] The mechanism used bears some resemblance to the rho-dependent termination in prokaryotes.[33] Eukaryotic cells contain hundreds of ribosomal DNA repeats, sometimes distributed over multiple chromosomes. Termination of transcription occurs in the ribosomal intergenic spacer region that contains several transcription termination sites upstream of a Pol I pausing site. Through a yet unknown mechanism, the 3’-end of the transcript is cleaved, generating a large primary rRNA molecule that is further processed into the mature 18S, 5.8S and 28S rRNAs.
As Pol II reaches the end of a gene, two protein complexes carried by the CTD, CPSF (cleavage and polyadenylation specificity factor) and CSTF (cleavage stimulation factor), recognize the poly-A signal in the transcribed RNA.[32] Poly-A-bound CPSF and CSTF recruit other proteins to carry out RNA cleavage and then polyadenylation. Poly-A polymerase adds approximately 200 adenines to the cleaved 3’ end of the RNA without a template.[32] The long poly-A tail is unique to transcripts made by Pol II.
In the process of terminating transcription by Pol I and Pol II, the elongation complex does not dissolve immediately after the RNA is cleaved. The polymerase continues to move along the template, generating a second RNA molecule associated with the elongation complex.[1] Two models have been proposed to explain how termination is achieved at last.[32] The allosteric model states that when transcription proceeds through the termination sequence, it causes disassembly of elongation factors and/or an assembly of termination factors that cause conformational changes of the elongation complex.[33][34] The torpedo model suggests that a 5' to 3' exonuclease degrades the second RNA as it emerges from the elongation complex. Polymerase is released as the highly processive exonuclease overtakes it. It is proposed that an emerging view will express a merge of these two models.[34]
Factor-independent[edit]
RNA polymerase III can terminate transcription efficiently without the involvement of additional factors. The Pol III termination signal consists of a stretch of thymines (on the nontemplate strand) located within 40bp downstream from the 3' end of mature RNAs.[32] The poly-T termination signal pauses Pol III and causes it to backtrack to the nearest RNA hairpin to become a “dead-end” complex.[35] Consistent with the allosteric mechanism of termination,[36] the RNA hairpin allosterically opens Pol III and causes the elongation complex to disintegrate. The extensive structure embedded in the Pol III-transcript thus is responsible for the factor-independent release of Pol III at the end of a gene. RNA-duplex-dependent termination is an ancient mechanism that dates back to the last universal common ancestor.
Eukaryotic transcriptional control[edit]
The regulation of gene expression in eukaryotes is achieved through the interaction of several levels of control that acts both locally to turn on or off individual genes in response to a specific cellular need and globally to maintain a chromatin-wide gene expression pattern that shapes cell identity.[1][37] Because eukaryotic genome is wrapped around histones to form nucelosomes and higher-order chromatin structures, the substrates for transcriptional machinery are in general partially concealed.[1] Without regulatory proteins, many genes are expressed at low level or not expressed at all. Transcription requires displacement of the positioned nucleosomes to enable the transcriptional machinery to gain access of the DNA.[38]
All steps in the transcription are subject to some degree of regulation.[1] Transcription initiation in particular is the primary level at which gene expression is regulated. Targeting the rate-limiting initial step is the most efficient in terms of energy costs for the cell. Transcription initiation is regulated by cis-acting elements (enhancers, silencers, isolators) within the regulatory regions of the DNA, and sequence-specific trans-acting factors that act as activators or repressors.[1] Gene transcription can also be regulated post-initiation by targeting the movement of the elongating polymerase.[39]
Global control and epigenetic regulation[edit]
Main article: Epigenetic regulation
The eukaryotic genome is organized into a compact chromatin structure that allows only regulated access to DNA. The chromatin structure can be globally "open" and more transcriptionally permissive, or globally "condensed" and transcriptionally inactive. The former (euchromatin) is lightly packed and rich in genes under active transcription. The latter (heterochromatin) includes gene-poor regions such as telomeres and centromeres but also regions with normal gene density but transcriptionally silenced. Transcription can be silenced by histone modification (deaceltylation and methylation), RNA interference, and/or DNA methylation.[40]
The gene expression patterns that define cell identity are inherited through cell division.[1] This process is called epigenetic regulation. DNA methylation is reliably inherited through the action of maintenance methylases that modify the nascent DNA strand generated by replication.[1] In mammalian cells, DNA methylation is the primary marker of transcriptionally silenced regions. Specialized proteins can recognize the marker and recruit histone deacetylases and methylases to re-establish the silencing. Nucleosome histone modifications could also be inherited during cell division, however, it is not clear whether it can work independently without the direction by DNA methylation.[1]
Gene-specific activation[edit]
The two main tasks of transcription initiation are to provide RNA polymerase with an access to the promoter and to assemble general transcription factors with polymerase into a transcription initiation complex. Diverse mechanisms of initiating transcription by overriding inhibitory signals at the gene promoter have been identified.[1] Eukaryotic genes have acquired extensive regulatory sequences that encompass a large number of regulator-binding sites and spread overall kilobases (sometimes hundreds of kilobases) from the promoter–-both upstream and downstream.[1] The regulator binding sites are often clustered together into units called enhancers. Enhancers can facilitate highly cooperative action of several transcription factors (which constitute enhanceosomes). Remote enhancers allow transcription regulation at a distance. Insulators situated between enhancers and promoters help define the genes that an enhancer can or cannot influence.
Eukaryotic transcriptional activators have separate DNA-binding and activating functions.[1] Upon binding to its cis-element, an activator can recruit polymerase directly or recruit other factors needed by the transcriptional machinery. An activator can also recruit nucleosome modifiers that alter chromatin in the vicinity of the promoter and thereby help initiation. Multiple activators can work together, either by recruiting a common or two mutually dependent components of the transcriptional machinery, or by helping each other bind to their DNA sites.[1] These interactions can synergize multiple signaling inputs and produce intricate transcriptional responses to address cellular needs.
Gene-specific repression[edit]
Eukaryotic transcription repressors share some of the mechanisms used by their prokaryotic counterparts. For example, by binding to a site on DNA that overlaps with the binding site of an activator, a repressor can inhibit binding of the activator. But more frequently, eukaryotic repressors inhibit the function of an activator by masking its activating domain, preventing its nuclear localization, promoting its degradation, or inactivating it through chemical modifications.[1] Repressors can directly inhibit transcription initiation by binding to a site upstream of a promoter and interacting with the transcriptional machinery. Repressors can indirectly repress transcription by recruiting histone modifiers (deacetylases and methylases) or nucelosome remodeling enzymes that affect the accessibility of the DNA.[1] Repressing histone and DNA modifications are also the basis of transcriptional silencing that can spread along the chromatin and switch off multiple genes.[41]
Elongation and termination control[edit]
The elongation phase starts once assembly of the elongation complex has been completed, and progresses until a termination sequence is encountered.[1] The post-initiation movement of RNA polymerase is the target of another class of important regulatory mechanisms. For example, the transcriptional activator Tat affects elongation rather than initiation during its regulation of HIV transcription.[42] In fact, many eukaryotic genes are regulated by releasing a block to transcription elongation called promoter-proximal pausing.[43] Pausing can influence chromatin structure at promoters to facilitate gene activity and lead to rapid or synchronous transcriptional responses when cells are exposed to an activation signal.[30] Pausing is associated with the binding of two negative elongation factors, DSIF (SPT4/SPT5) and NELF, to the elongation complex. Other factors can also influence the stability and duration of the paused polymerase.[44] Pause release is triggered by the recruitment of the P-TEFb kinase.[39]
Transcription termination has also emerged as an important area of transcriptional regulation. Termination is coupled with the efficient recycling of polymerase.[45] The factors associated with transcription termination can also mediate gene looping and thereby determine the efficiency of re-initiation.
Transcription-coupled DNA repair[edit]
When transcription is arrested by the presence of a lesion in the transcribed strand of a gene, DNA repair proteins are recruited to the stalled RNA polymerase to initiate a process called transcription-coupled repair.[46] Central to this process is the general transcription factor TFIIH that has ATPase activity. TFIIH causes a conformational change in the polymerase, to expose the transcription bubble trapped inside, in order for the DNA repair enzymes to gain access to the lesion.[47] Thus, RNA polymerase serves as damage-sensing protein in the cell to target repair enzymes to genes that are being actively transcribed.
Comparisons between prokaryotic and eukaryotic transcription[edit]
Eukaryotic transcription is more complex than prokaryotic transcription. For instance, in eukaryotes the genetic material (DNA), and therefore transcription, is primarily localized to the nucleus, where it is separated from the cytoplasm (in which translation occurs) by the nuclear membrane. This allows for the temporal regulation of gene expression through the sequestration of the RNA in the nucleus, and allows for selective transport of mature RNAs to the cytoplasm. Bacteria do not have a distinct nucleus that separates DNA from ribosome and mRNA is translated into protein as soon as it is transcribed. The coupling between the two processes provides an important mechanism for prokaryotic gene regulation.[1]
At the level of initiation, RNA polymerase in prokaryotes (bacteria in particular) binds strongly to the promoter region and initiates a high basal rate of transcription. No ATP hydrolysis is needed for the close-to-open transition, promoter melting is driven by binding reactions that favor the melted conformation. Chromatin greatly impedes transcription in eukaryotes. Assembly of large multi-protein preinitiation complex is required for promoter-specific initiation. Promoter melting in eukaryotes requires hydrolysis of ATP. As a result, eukaryotic RNA polymerases exhibit a low basal rate of transcription initiation.[41]
Regulation of transcription in cancer[edit]
Main article: Regulation of transcription in cancer
In vertebrates, the majority of gene promoters contain a CpG island with numerous CpG sites.[48] When many of a gene's promoter CpG sites are methylated the gene becomes silenced.[49] Colorectal cancers typically have 3 to 6 driver mutations and 33 to 66 hitchhiker or passenger mutations.[50] However, transcriptional silencing may be of more importance than mutation in causing progression to cancer. For example, in colorectal cancers about 600 to 800 genes are transcriptionally silenced by CpG island methylation (see regulation of transcription in cancer). Transcriptional repression in cancer can also occur by other epigenetic mechanisms, such as altered expression of microRNAs.[51] In breast cancer, transcriptional repression of BRCA1 may occur more frequently by over-expressed microRNA-182 than by hypermethylation of the BRCA1 promoter (see Low expression of BRCA1 in breast and ovarian cancers).
References[edit]
^ Jump up to: a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj Watson, James; Tania A. Baker; Stephen P. Bell; Alexander Gann; Michael Levine; Richard Losik; Stephen C. Harrison. Molecular Biology of the Gene (7th ed.). Benjamin-Cummings Publishing Company. ISBN 978-0-321-76243-6.
^ Jump up to: a b Mattick, J. S. (1 November 2001). "Non-coding RNAs: the architects of eukaryotic complexity". EMBO Reports. 2 (11): 986–991. doi:10.1093/embo-reports/kve230. PMC 1084129  . PMID 11713189.
^ Jump up to: a b c d Lodish Harvey, ...[et al]. Molecular cell biology (7th ed.). New York: W.H. Freeman and Co. ISBN 9781429234139.
^ Jump up to: a b c Cramer P, Armache KJ, Baumli S, Benkert S, Brueckner F, Buchen C, Damsma GE, Dengl S, Geiger SR, Jasiak AJ, Jawhari A, Jennebach S, Kamenski T, Kettenberger H, Kuhn CD, Lehmann E, Leike K, Sydow JF, Vannini A (2008). "Structure of eukaryotic RNA polymerases". Annu Rev Biophys. 37: 337–52. doi:10.1146/annurev.biophys.37.032807.130008. PMID 18573085.
Jump up ^ Sirri, Valentina; Silvio Urcuqui-Inchima; Pascal Roussel; Danièle Hernandez-Verdun (2008). "Nucleolus: the fascinating nuclear body". Histochem Cell Biol. 129 (1): 13–31. doi:10.1007/s00418-007-0359-6. PMC 2137947  . PMID 18046571.
Jump up ^ Fromont-Racine, Micheline; Senger, Bruno; Saveanu, Cosmin; Fasiolo, Franco (August 2003). "Ribosome assembly in eukaryotes". Gene. 313: 17–42. doi:10.1016/S0378-1119(03)00629-2.
Jump up ^ Dieci, Giorgio; Fiorino, Gloria; Castelnuovo, Manuele; Teichmann, Martin; Pagano, Aldo (December 2007). "The expanding RNA polymerase III transcriptome". Trends in Genetics. 23 (12): 614–622. doi:10.1016/j.tig.2007.09.001. PMID 17977614.
^ Jump up to: a b Carter, R.; Drouin, G. (21 December 2009). "The Increase in the Number of Subunits in Eukaryotic RNA Polymerase III Relative to RNA Polymerase II Is due to the Permanent Recruitment of General Transcription Factors". Molecular Biology and Evolution. 27 (5): 1035–1043. doi:10.1093/molbev/msp316. PMID 20026480.
Jump up ^ Fernández-Tornero, Carlos; Moreno-Morcillo, María; Rashid, Umar J.; Taylor, Nicholas M. I.; Ruiz, Federico M.; Gruene, Tim; Legrand, Pierre; Steuerwald, Ulrich; Müller, Christoph W. (23 October 2013). "Crystal structure of the 14-subunit RNA polymerase I". Nature. 502 (7473): 644–649. doi:10.1038/nature12636.
Jump up ^ Cramer, P. (19 April 2001). "Structural Basis of Transcription: RNA Polymerase II at 2.8 Angstrom Resolution". Science. 292 (5523): 1863–1876. doi:10.1126/science.1059493. PMID 11313498.
^ Jump up to: a b Corden, Jeffry L. (13 November 2013). "RNA Polymerase II C-Terminal Domain: Tethering Transcription to Transcript and Template". Chemical Reviews. 113 (11): 8423–8455. doi:10.1021/cr400158h. PMC 3988834  . PMID 24040939.
Jump up ^ Butler, Jennifer; James T. Kadonaga (2002). "The RNA polymerase II core promoter: a key component in the regulation of gene expression". Genes Dev. 16 (20): 2583–2592. doi:10.1101/gad.1026202. PMID 12381658.
Jump up ^ Lenhard, Boris; Sandelin, Albin; Carninci, Piero (6 March 2012). "Metazoan promoters: emerging characteristics and insights into transcriptional regulation". Nature Reviews Genetics. 13: 233–45. doi:10.1038/nrg3163. PMID 22392219.
Jump up ^ Orphanides, G; Lagrange, T; Reinberg, D (1 November 1996). "The general transcription factors of RNA polymerase II". Genes & Development. 10 (21): 2657–2683. doi:10.1101/gad.10.21.2657. PMID 8946909.
Jump up ^ ROEDER, R (September 1996). "The role of general initiation factors in transcription by RNA polymerase II". Trends in Biochemical Sciences. 21 (9): 327–335. doi:10.1016/S0968-0004(96)10050-5.
Jump up ^ He, Yuan; Jie Fang; Dylan J. Taatjes; Eva Nogales (28 March 2013). "Structural visualization of key steps in human transcription initiation". Nature. 495: 481–486. doi:10.1038/nature11991. PMC 3612373  . PMID 23446344.
^ Jump up to: a b Holstege, F. C.P. (15 December 1997). "Three transitions in the RNA polymerase II transcription complex during initiation". The EMBO Journal. 16 (24): 7468–7480. doi:10.1093/emboj/16.24.7468. PMC 1170346  . PMID 9405375.
Jump up ^ SVEJSTRUP, J; VICHI, P; EGLY, J (September 1996). "The multiple roles of transcription/repair factor TFIIH". Trends in Biochemical Sciences. 21 (9): 346–350. doi:10.1016/S0968-0004(96)10046-3. PMID 8870499.
^ Jump up to: a b Revyakin A, Liu C, Ebright RH, Strick TR (17 November 2006). "Abortive Initiation and Productive Initiation by RNA Polymerase Involve DNA Scrunching". Science. 314 (5802): 1139–1143. doi:10.1126/science.1131398. PMC 2754787  . PMID 17110577.
Jump up ^ Dvir, Arik (September 2002). "Promoter escape by RNA polymerase II". Biochimica et Biophysica Acta (BBA) - Gene Structure and Expression. 1577 (2): 208–223. doi:10.1016/S0167-4781(02)00453-0. PMID 12213653.
Jump up ^ Pokholok, Dmitry K; Hannett, Nancy M; Young, Richard A (April 2002). "Exchange of RNA Polymerase II Initiation and Elongation Factors during Gene Expression In Vivo". Molecular Cell. 9 (4): 799–809. doi:10.1016/S1097-2765(02)00502-6.
Jump up ^ Wade, Joseph T; Struhl, Kevin (April 2008). "The transition from transcriptional initiation to elongation". Current Opinion in Genetics & Development. 18 (2): 130–136. doi:10.1016/j.gde.2007.12.008.
Jump up ^ Saunders, Abbie; Core, Leighton J.; Lis, John T. (August 2006). "Breaking barriers to transcription elongation". Nature Reviews Molecular Cell Biology. 7 (8): 557–567. doi:10.1038/nrm1981. PMID 16936696.
Jump up ^ Arndt, Karen; Caroline Kane (1 October 2003). "Running with RNA polymerase: eukaryotic transcript elongation". Trends in Genetics. 19 (10): 543–550. doi:10.1016/j.tig.2003.08.008. PMID 14550628.
^ Jump up to: a b Brès, Vanessa; Yoh, Sunnie M; Jones, Katherine A (June 2008). "The multi-tasking P-TEFb complex". Current Opinion in Cell Biology. 20 (3): 334–340. doi:10.1016/j.ceb.2008.04.008.
Jump up ^ Westover, Kenneth D.; Bushnell, David A.; Kornberg, Roger D. (November 2004). "Structural Basis of Transcription". Cell. 119 (4): 481–489. doi:10.1016/j.cell.2004.10.016.
Jump up ^ Wang, Dong; Bushnell, David A.; Westover, Kenneth D.; Kaplan, Craig D.; Kornberg, Roger D. (December 2006). "Structural Basis of Transcription: Role of the Trigger Loop in Substrate Specificity and Catalysis". Cell. 127 (5): 941–954. doi:10.1016/j.cell.2006.11.023. PMC 1876690  . PMID 17129781.
Jump up ^ Wang, D.; Bushnell, D. A.; Huang, X.; Westover, K. D.; Levitt, M.; Kornberg, R. D. (28 May 2009). "Structural Basis of Transcription: Backtracked RNA Polymerase II at 3.4 Angstrom Resolution". Science. 324 (5931): 1203–1206. doi:10.1126/science.1168729. PMC 2718261  . PMID 19478184.
Jump up ^ Sosunov, Vasily; Ekaterina Sosunova; Arkady Mustaev; Irina Bass; Vadim Nikiforov; Alex Goldfarb (2003). "Unified two-metal mechanism of RNA synthesis and degradation by RNA polymerase". The EMBO Journal. 22: 2234–2244. doi:10.1093/emboj/cdg193. PMC 156065  . PMID 12727889.
^ Jump up to: a b c Adelman, Karen; Lis, John T. (18 September 2012). "Promoter-proximal pausing of RNA polymerase II: emerging roles in metazoans". Nature Reviews Genetics. 13 (10): 720–731. doi:10.1038/nrg3293.
Jump up ^ Missra, A.; Gilmour, D. S. (4 June 2010). "Interactions between DSIF (DRB sensitivity inducing factor), NELF (negative elongation factor), and the Drosophila RNA polymerase II transcription elongation complex". Proceedings of the National Academy of Sciences. 107 (25): 11301–11306. doi:10.1073/pnas.1000681107. PMC 2895096  . PMID 20534440.
^ Jump up to: a b c d e Richard, P.; Manley, J. L. (1 June 2009). "Transcription termination by nuclear RNA polymerases". Genes & Development. 23 (11): 1247–1269. doi:10.1101/gad.1792809.
^ Jump up to: a b Clancey, Suzanne (2008). "DNA transcription". Nature Education. 1 (41). Retrieved 27 November 2013.
^ Jump up to: a b Rosonina, E. (1 May 2006). "Terminating the transcript: breaking up is hard to do". Genes & Development. 20 (9): 1050–1056. doi:10.1101/gad.1431606. PMID 16651651.
Jump up ^ Nielsen, S.; Yuzenkova, Y.; Zenkin, N. (27 June 2013). "Mechanism of Eukaryotic RNA Polymerase III Transcription Termination". Science. 340 (6140): 1577–1580. doi:10.1126/science.1237934.
Jump up ^ Epshtein, Vitaly; Cardinale, Christopher J.; Ruckenstein, Andrei E.; Borukhov, Sergei; Nudler, Evgeny (December 2007). "An Allosteric Path to Transcription Termination". Molecular Cell. 28 (6): 991–1001. doi:10.1016/j.molcel.2007.10.011.
Jump up ^ Shandilya, J; Robert SG (2012). "The transcription cycle in eukaryotes: from productive initiation to RNA polymerase II recycling". Biochim Biophys Acta. 1819 (5): 391–400. doi:10.1016/j.bbagrm.2012.01.010. PMID 22306664.
Jump up ^ Kulaeva, Olga; Daria Gaykalova; Vasily M. Studitsky (2007). "Transcription Through Chromatin by RNA polymerase II: Histone Displacement and Exchange". Mutat Res. 618 (1–2): 116–129. doi:10.1016/j.mrfmmm.2006.05.040. PMC 1924643  . PMID 17313961.
^ Jump up to: a b Peterlin, BM; DH Price (2006). "Controlling the elongation phase of transcription with P-TEFb". Molecular Cell. 23 (3): 297–305. doi:10.1016/j.molcel.2006.06.014. PMID 16885020.
Jump up ^ Bannister, Andrew J.; Kouzarides, Tony (15 February 2011). "Regulation of chromatin by histone modifications". Cell Research. 21 (3): 381–395. doi:10.1038/cr.2011.22. PMC 3193420  . PMID 21321607.
^ Jump up to: a b Brown, Terrance A. (2002). Genomes. New York: Wiley-Liss. ISBN 0-471-25046-5.
Jump up ^ Kao, Shaw-Yi; Andrew Calman; Paul Luciw; Matija Peterlin (3 December 1987). "Anti-termination of transcription within the long terminal repeat of HIV-1 by tat gene product". Nature. 330 (6147): 489–493. doi:10.1038/330489a0. PMID 2825027.
Jump up ^ Lis, J (1998). "Promoter-associated Pausing in Promoter Architecture and Postinitiation Transcriptional Regulation". Cold Spring Harb Symp Quant Biol. 63: 347–56. doi:10.1101/sqb.1998.63.347. PMID 10384299.
Jump up ^ Cheng B, Li T, Rahl PB, Adamson TE, Loudas NB, Guo J, Varzavand K, Cooper JJ, Hu X, Gnatt A, Young RA, Price DH (January 2012). "Functional Association of Gdown1 with RNA Polymerase II Poised on Human Genes". Molecular Cell. 45 (1): 38–50. doi:10.1016/j.molcel.2011.10.022. PMC 3259526  . PMID 22244331.
Jump up ^ Feige, MJ; LM Hendershot (April 2011). "Disulfide bonds in ER protein folding and homeostasis". Curr Opin Cell Biol. 23 (2): 167–75. doi:10.1016/j.ceb.2010.10.012. PMC 3078216  . PMID 21144725.
Jump up ^ MELLON, I (October 1987). "Selective removal of transcription-blocking DNA damage from the transcribed strand of the mammalian DHFR gene". Cell. 51 (2): 241–249. doi:10.1016/0092-8674(87)90151-6.
Jump up ^ Sarker, Altaf H.; Tsutakawa, Susan E.; Kostek, Seth; Ng, Cliff; Shin, David S.; Peris, Marian; Campeau, Eric; Tainer, John A.; Nogales, Eva; Cooper, Priscilla K. (October 2005). "Recognition of RNA Polymerase II and Transcription Bubbles by XPG, CSB, and TFIIH: Insights for Transcription-Coupled Repair and Cockayne Syndrome". Molecular Cell. 20 (2): 187–198. doi:10.1016/j.molcel.2005.09.022. PMID 16246722.
Jump up ^ Saxonov S, Berg P, Brutlag DL (2006). "A genome-wide analysis of CpG dinucleotides in the human genome distinguishes two distinct classes of promoters". Proc. Natl. Acad. Sci. U.S.A. 103 (5): 1412–7. doi:10.1073/pnas.0510310103. PMC 1345710  . PMID 16432200.
Jump up ^ Bird A (2002). "DNA methylation patterns and epigenetic memory". Genes Dev. 16 (1): 6–21. doi:10.1101/gad.947102. PMID 11782440.
Jump up ^ Vogelstein B, Papadopoulos N, Velculescu VE, Zhou S, Diaz LA, Kinzler KW (2013). "Cancer genome landscapes". Science. 339 (6127): 1546–58. doi:10.1126/science.1235122. PMC 3749880  . PMID 23539594.
Jump up ^ Tessitore A, Cicciarelli G, Del Vecchio F, Gaggiano A, Verzella D, Fischietti M, Vecchiotti D, Capece D, Zazzeroni F, Alesse E (2014). "MicroRNAs in the DNA Damage/Repair Network and Cancer". Int J Genomics. 2014: 820248. doi:10.1155/2014/820248. PMC 3926391  . PMID 24616890.
[show]
v
t
e
Gene expression
Introduction
to genetics
Genetic code
Central dogma
DNA → RNA → Protein
Special transfers
RNA→RNA
RNA→DNA
Protein→Protein
Transcription
Types
Bacterial
Eukaryotic
Key elements
Transcription factor
RNA polymerase
Promoter
Post-transcription
Precursor mRNA (pre-mRNA / hnRNA)
5' capping
Splicing
Polyadenylation
Histone acetylation and deacetylation
Translation
Types
Prokaryotic
Eukaryotic
Key elements
Ribosome
Transfer RNA (tRNA)
Ribosome-nascent chain complex (RNC)
Post-translational modification (functional groups · peptides · structural changes)
Regulation
Epigenetic
imprinting
Transcriptional
Gene regulatory network
cis-regulatory element
lac operon
Post-transcriptional
sequestration (P-bodies)
alternative splicing
microRNA
Translational
Post-translational
reversible
irreversible
Influential people
François Jacob
Jacques Monod
[show]
v
t
e
Transcription (Bacterial, Eukaryotic)
Transcriptional regulation
prokaryotic
Operon
lac operon
trp operon
gab operon
ara operon
gal operon
Repressor
lac repressor
trp repressor
eukaryotic
Histone-modifying enzymes
(histone/nucleosome):
Histone methylation/Histone methyltransferase
EZH2
Histone demethylase
Histone acetylation and deacetylation
Histone deacetylase HDAC1
Histone acetyltransferase
DNA methylation:
DNA methyltransferase
Chromatin remodeling:
CHD7
both
Transcription coregulator
Coactivator
Corepressor
Inducer
Promotion
Promoter
Pribnow box
TATA box
BRE
CAAT box
Response element
Enhancer
E-box
Response element
Insulator
Silencer
Internal control region
Initiation (bacterial,
eukaryotic
Transcription start site
Elongation
bacterial RNA polymerase: rpoB
eukaryotic RNA polymerase: RNA polymerase II
Termination
(bacterial,
eukaryotic)
Terminator
Intrinsic termination
Rho factor
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