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Hilton Worldwide
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Hilton Worldwide Holdings Inc.
Headquarters in Tysons Corner, Virginia, United States
Formerly called
Hilton Hotels Corporation
Type
Public
Traded as
NYSE: HLT
S&P 500 component
Industry
Hospitality
Founded
May 31, 1919; 98 years ago (1919-05-31) in Cisco, Texas, United States
(as Hilton Hotels Corporation)
Founder
Conrad Hilton
Headquarters
Tysons Corner, Virginia, United States
Number of locations
5,284 (December 2017)[1][2]
Area served
Worldwide
Key people
Christopher J. Nassetta (President and CEO)
Kevin J. Jacobs (CFO)
Revenue
US$9.140 billion (2017)[1]
Operating income
US$1.372 billion (2017)[1]
Net income
US$1.259 billion (2017)[1]
Total assets
US$14.308 billion (2017)[1]
Total equity
US$2.072 billion (2017)[1]
Number of employees
~163,000 (December 2017)[1]
Website
hilton.com
Hilton Worldwide Holdings Inc., formerly Hilton Hotels Corporation, is an American multinational hospitality company that manages and franchises a broad portfolio of hotels and resorts. Founded by Conrad Hilton in 1919, the corporation is now led by Christopher J. Nassetta.
Hilton is headquartered in Tysons Corner, Virginia. As of December 2017, its portfolio includes more than 5,200 properties (including timeshare properties) with over 856,000 rooms in 105 countries and territories.[3] Prior to their December 2013 IPO, Hilton was ranked as the 36th largest privately held company in the United States by Forbes.[4]
Hilton has 14 brands across different market segments, including Conrad Hotels & Resorts, Canopy by Hilton, Curio - A Collection by Hilton, Hilton Hotels & Resorts, DoubleTree by Hilton, Embassy Suites Hotels, Hilton Garden Inn, Hampton by Hilton, Homewood Suites by Hilton, Home2 Suites by Hilton, Hilton Grand Vacations and Waldorf Astoria Hotels & Resorts, Tru by Hilton, and Tapestry Collection by Hilton.
On December 12, 2013, Hilton again became a public company, raising an estimated $2.35 billion in its second IPO. At the time, The Blackstone Group held a 45.8 percent stake in the company.[5] In October 2016, HNA Group agreed to acquire a 25 percent equity interest in Hilton from Blackstone. The transaction was expected to close in the first quarter of 2017.[6][7][8] Hilton's largest stockholders are HNA Group, Blackstone, and Wellington Management Group, which own 25%, 15.2%, and 6.7% of Hilton common stock respectively.[9]
Hilton was founded by Conrad Hilton in Cisco, Texas, in 1919 and had its headquarters in Beverly Hills, California, from 1969 until 2009. In August 2009, the company moved to Tysons Corner, unincorporated Fairfax County, Virginia, near McLean.[10][11][12]
Contents
[hide]
1 History
1.1 Two chains with one name
1.2 21st century
2 Brands
2.1 Luxury
2.2 Full service
2.3 Focused service
2.4 All suites
2.5 Lifestyle
2.6 Timeshare
2.7 Former
2.8 Franchising
2.9 Hilton Honors
3 Corporate affairs
3.1 Company culture
4 Hilton in popular culture
5 References
History[edit]
To be the preeminent global hospitality company - the first choice of guests, team members, and owners alike.
“
”
Hilton's mission[when?][13]
In 1919, Conrad Hilton purchased his first hotel, the 40-room Mobley Hotel in Cisco, Texas, and bought additional Texas hotels as years passed.[14]
In 1925, the Dallas Hilton became the first hotel to use the Hilton name.[14] In 1927, Hilton expanded to Waco, Texas,[15] where he opened the first hotel with air-conditioning in public areas and cold running water.[16][17][18]
In 1943, Hilton purchased the Roosevelt Hotel and the Plaza Hotel in New York, establishing the first hospitality company to span the contiguous United States.[19] The company incorporated in 1946 as the Hilton Hotels Corporation, and subsequently began public trading of shares on the New York Stock Exchange.[20][21][22]
The entrance to the Waldorf-Astoria in New York, Conrad's "greatest of them all"
Hilton International was founded a few years later, in 1949, with the opening of the Caribe Hilton Hotel in Puerto Rico.[23] Barman Ramon "Monchito" Marreno claimed he created the piña colada cocktail at this resort.[24][25] Hilton purchased the Waldorf Astoria New York in the same year.[26][27]
The Hotels Statler Company was acquired in 1954 for $111 million in what was then the world's most expensive real estate transaction.[28] One year later, Hilton created the world's first central reservations office, titled "HILCRON". The reservations team in 1955 consisted of eight members on staff booking reservations for any of Hilton's then 28 hotels. Reservations agents used the "availability board" to track records. The chalk board measured 30 feet by 6 feet, and allowed HILCRON to make over 6,000 reservations in the year.[29] Bookings could be made for any Hilton via telephone, telegram, or Teletype.
Later in 1955, Hilton launched a program to ensure every hotel room would include air conditioning.[30]
In late 1955, Hilton opened the first post-World War II property in Istanbul.[30][31]
Hilton is credited with pioneering the airport hotel concept with the opening of the San Francisco Airport Hilton in 1959.[32]
In 1965, Hilton launched Lady Hilton, the first hotel concept created specifically for women guests.[33] To appeal to female travelers, a number of properties offered floors occupied by only women along with distinct amenities for their usage.[34][35]
In 1969, the first DoubleTree Hotel opened. However, Hilton was not affiliated with the brand until its acquisition of the parent company in 1999.[36][37]
Hilton purchased the Flamingo Las Vegas in 1970,[38][39] which would become the first in the domestic gaming business to be listed on the NYSE.[40]
In 1979, founder Conrad Hilton died at the age of 91.[41] Hilton Hotels Corporation later created the Conrad Hotels brand in honor of Hilton.[42]
Hilton Honors (formerly Hilton HHonors), the company's guest loyalty program, was initiated in 1987.[43] In 1994, the Honors surpassed competing hotel loyalty programs by offering members both hotel credit points and airline credit miles.[44][45]
The company has been a sponsor of the United States Olympic Team.[46][47]
Two chains with one name[edit]
The company spun off its international operations into a separately traded company on December 1, 1964, known as "Hilton International Co." It was acquired in 1967 by Trans World Corp., the holding company for Trans World Airlines. In 1986, it was sold to UAL Corp., the holding company for United Airlines, which became Allegis Corp. in an attempt to re-incarnate itself as a full-service travel company, encompassing Westin Hotels and Hertz rental cars in addition to Hilton International and United Airlines. In 1987, after a corporate putsch, the renamed UAL Corp. sold Hilton International to Ladbroke Group plc, a British leisure and gambling company, which, in May 1999, adopted the name "Hilton Group plc."[48]
The former Hilton Hotels Corporation headquarters in Beverly Hills
As a result, there were two separate, fully independent companies operating hotels under the Hilton name. Those Hilton Hotels outside the U.S. were, until 2006, styled as "Hilton International" hotels. Because the two chains were contractually forbidden to operate hotels in the other's territory under the Hilton name, for many years hotels run by Hilton International in the U.S. were called Vista International Hotels, while hotels operated by the American arm of Hilton outside the U.S. were named Conrad Hotels.[49]
In 1997, to minimize longtime consumer confusion, the American and British Hilton companies adopted a joint marketing agreement under which they shared the same logos, promoted each other's brands and maintained joint reservation systems. At that point, the Vista chain was phased out, while Conrad has been restyled as one of the luxury brands of Hilton (along with the Waldorf-Astoria Collection) and operates hotels within the US as well as abroad.
In 1971, Hilton acquired International Leisure Company, including the Las Vegas Hilton and Flamingo Hilton.[50]
In 1998, Hilton spun off its gaming operations into a separate, publicly held company called Park Place Entertainment (later Caesars Entertainment, Inc.)[citation needed]
In 1999, Hilton acquired Promus Hotel Corporation, which included the DoubleTree, Red Lion, Embassy Suites, Hampton Inn and Homewood Suites brands.[51]
21st century[edit]
Hilton Hotels & Resorts, Hilton's flagship brand
In 2001, Hilton agreed to sell Red Lion to WestCoast Hospitality.[52]
On December 29, 2005, Hilton Hotels Corporation agreed to re-acquire the Hilton International chain from its British owner, Hilton Group plc, for £3.3 billion ($5.71 billion). As well as bringing the two Hilton companies back together as a single entity, this deal also included Hilton plc properties operating as Conrad Hotels, Scandic Hotels and LivingWell Health Clubs.[53] On February 23, 2006, the deal closed, making Hilton Hotels the world's fifth largest hotel operator in number of rooms.[54] Hilton Group PLC (headquartered in the UK) then renamed itself Ladbrokes plc. Scandic Hotels was sold the next year on March 1 to EQT Partners.[55]
Christopher J. Nassetta in 2004
On July 3, 2007, Hilton Hotels Corp. agreed to an all-cash buyout from the Blackstone Group LP in a $26 billion (including debt) deal that would make Blackstone the world's largest hotel owner.[56] At $47.50 per share, the buyout price was 32 percent higher than the closing value of a share of Hilton stock on July 3.[57] The deal was the culmination of a year of on and off discussions with Blackstone.[58] In October 2007, Christopher J. Nassetta was appointed President and Chief Executive Officer of Hilton.[59] In February 2009, Hilton Hotels Corp. announced that its headquarters were moving from Beverly Hills to Fairfax County, Virginia.[60]
While Blackstone saw an opportunity to streamline the company and push Hilton's expansion overseas when Blackstone pursued Hilton in 2006 and 2007, the buyout saddled the company with $20 billion of debt just as the economy was turning down. The debt had very liberal terms, so there was no danger of default, but when travel slowed, the company suffered. In April 2010, Hilton and Blackstone restructured the debt. Blackstone invested a further $800 million of equity and the debt was reduced to $16 billion.[61]
Hilton returned to being a public company on December 12, 2013. This second IPO in the company's history raised an estimated $2.35 billion.[62] The Blackstone Group retained a 45.8% stake in the company.[5]
The company announced in February 2016 that Hilton would turn its hotel holdings into a real estate investment trust. Prior to making the announcement, the company went to the IRS for approval.[63]
In February 2016, Hilton announced its intention to spin off its timeshare and real estate businesses, creating three independent public companies.[64]
Brands[edit]
Hilton San Diego Bayfront, one of Hilton's flagship properties
Luxury[edit]
Conrad Hotels & Resorts
Waldorf Astoria Hotels and Resorts
Full service[edit]
Curio Collection by Hilton
Hilton Hotels & Resorts (Flagship Brand of Hilton Worldwide)
Tapestry Collection by Hilton
DoubleTree by Hilton
Homewood Suites by Hilton entrance sign
Focused service[edit]
Hampton by Hilton
Hilton Garden Inn
Tru by Hilton
All suites[edit]
Embassy Suites by Hilton
Home2 Suites by Hilton
Homewood Suites by Hilton
Lifestyle[edit]
Canopy by Hilton[65]
Timeshare[edit]
Hilton Grand Vacations
Former[edit]
Coral by Hilton
Denizen Hotels
Lady Hilton
Scandic Hotels
Franchising[edit]
As of 2013, about 70% of the rooms branded under Hilton were franchised to independent operators and companies. During its 2007–2013 ownership of Hilton, Blackstone Group pursued a strategy of predominantly expanding Hilton's reach through franchise agreements, while relatively few new properties were actually operated by Hilton. Hence, the proportion of franchised rooms grew significantly during this period.[66] The practice of franchising is popular within the hospitality industry among most major hotel chains, as the parent company does not have to pay for the maintenance and overhead costs of franchised properties. Franchisees must follow strict brand standards in order to maintain a licensing agreement with Hilton.[66] Many of Hilton's flagship properties, airport properties, and largest resorts, however, are corporately managed.
Hilton Honors[edit]
Hilton Honors (formerly Hilton HHonors) is Hilton's guest loyalty program through which frequent guests can accumulate points and airline miles by staying within the Hilton portfolio. The program is one of the largest of its type, with approximately 71 million members.[67]
Hilton renamed the Hilton HHonors program to Hilton Honors in February 2017.[68]
Corporate affairs[edit]
The company has its headquarters in Tysons Corner, Virginia and an operations center in Memphis. Its Asia-Pacific operations are managed out of Singapore, its Middle East and Africa operations are managed out of Dubai, and its European operations are based in the UK.[69]
Company culture[edit]
Hilton San Francisco and Towers, the second largest Hilton with over 2,000 rooms
According to Careerbliss.com, Hilton ranked first in the list of "2012 Happiest Companies in America", with a score of 4.36 out of 5.[70] The survey looked at job reviews from more than 100,000 employees such as characteristics like work-life balance, company culture and reputation, and the relationships employees have with their bosses.[71] Hilton has scored 100% on the Corporate Equality Index each year from 2014 to 2017.[72]
In 2016, Hilton was named one of the "World's 25 Best Multinational Workplaces" by Fortune and Great Place to Work.[73][74][75]
In 2017, Fortune ranked Hilton number 26 in their list of "The 100 Best Companies to Work For".[76][77]
Hilton in popular culture[edit]
The 671-room Beverly Hilton in Beverly Hills, California is home to the Golden Globe Awards. It has played host to other awards shows, including the Soap Opera Awards.
Keith Richards and Mick Jagger performed in the East Penthouse on the 45th floor of the New York Hilton on October 28, 1965.[78]
John Lennon and Yoko Ono honeymooned in the presidential suite at the Hilton Amsterdam where they hosted one of their famous "Bed-Ins" for a full week in 1969.[79]
In 1971, Diamonds Are Forever was filmed at the Las Vegas Hilton.[80]
On April 3, 1973, Dr. Martin Cooper made the world's first cell phone call in front of the New York Hilton Midtown. A press conference was held at the hotel later that day to mark the milestone.[81][82]
The Bodyguard with Whitney Houston and Kevin Costner was filmed at the Fontainebleau Hilton in Miami in 1992.[83][84]
In 1995, the James Bond movie GoldenEye was filmed at the Langham Hilton.[85][86]
The Insider was filmed in 1999 at the Seelbach Hilton.[87]
In 2006, several movies including Spider-Man 3,[88] Michael Clayton and American Gangster were filmed at the New York Hilton.[89][90]
Conrad Hilton (played by actor Chelcie Ross) features as a major character in the third season of Mad Men, as lead character Don Draper creates a series of ad campaigns for Hilton Hotels.[91] The Drapers travel during one episode to the Cavalieri Hilton in Rome,[92] though the scenes were actually shot at the Dorothy Chandler Pavilion in Los Angeles.[93]
On the rotating wheel space station in Stanley Kubrick's film 2001: A Space Odyssey, a receptionist is shown sitting at the entrance to the Hilton Space Station 5.[94][95]
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[show]
v
t
e
Hilton Worldwide
Luxury & Lifestyle
Canopy by Hilton
Conrad Hotels & Resorts
Waldorf Astoria Hotels & Resorts
Full service
Curio Collection
DoubleTree by Hilton
Hilton Hotels & Resorts
Tapestry Collection by Hilton
All Suites
Embassy Suites by Hilton
Home2 Suites by Hilton
Homewood Suites by Hilton
Focused service
Hampton by Hilton
Hilton Garden Inn
Tru by Hilton
Vacation ownership
Hilton Grand Vacations Company
Loyalty program
Hilton Honors
Former divisions
Caesars Entertainment
Carte Blanche Card
Coral by Hilton
Denizen Hotels
Scandic Hotels
[show]
v
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Founders
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Stephen A. Schwarzman
Business Services
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Media/Telecommunications
SESAC
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Nielsen Holdings (minority stake)
Axtel (minority stake)
Deutsche Telekom (minority stake)
TDC (minority stake)
The Weather Channel (minority stake)
Travel & Leisure
Hilton Worldwide
Center Parcs Europe (freehold)
Caesars Entertainment Corporation (minority stake)
Merlin Entertainments (minority stake)
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[show]
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British Indian Association
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Bengal in 1857 was not prepared for systematic political efforts for the achievement of a well-defined political programme. There were, however, bold adventurers who could perceive dimly the inevitable trend in India’s political evolution. Harish Chandra Mukherjee wrote in the Hindu Patriot on 14 January 1858, in connection with the proposal for transferring the government of India to the Crown: “Can a revolution in the Indian government be authorised by Parliament without consulting the wishes of vast millions of men for whose benefit it is proposed to be made? The reply must be in the negative. The time has nearly come when all Indian questions must be solved by Indians.” [1]
Sengupta, Nitish
The British Indian Association was established on 31 October 1851. Its formation was a major event of 19th-century India. Its establishment meant Indians had come together and could no longer be ignored. It developed enormous hopes amongst the Indians about their future.[2]
The first committee of the association was composed of : Raja Radhakanta Deb – President, Raja Kalikrishna Deb – Vice-President, Debendranath Tagore – secretary, Digambar Mitra – Asst Secretary, members – Raja Staya Saran Ghosal, Harakumar Tagore, Prasanna Coomar Tagore, Ramanath Tagore, Jay Krishna Mukerjee, Asutosh Deb, Harimohan Sen, Ramgopal Ghosh, Umesh Chandra Dutta (Rambagan), Krishna Kishore Ghosh, Jagadananda Mukhopadhyay, Peary Chand Mitra, and Sambhunath Pandit.[3]
Most of the early leaders of the British Indian Association were conservatives by tradition and temperament, although there were some progressive like Ramgopal Ghosh and Peary Chand Mitra.[1]
It was formed by the amalgamation of the Landholders’ Society and the Bengal British India Society.[1]
Contents
[hide]
1 Landholders’ Society
2 British India Society
3 Objectives and extent of British Indian Association
4 Criticism
5 References
Landholders’ Society[edit]
It gave to the people the first lesson in the art of fighting constitutionally for their rights, and taught them manfully to assert their claims and give expression to their opinions. Ostensibly, it advocated the rights of zamindars, but as their rights were intimately bound up with those of the ryots, the one cannot be separated from the other.[1]
Mitra, Rajendra Lal
The Zamindari Association, which was later renamed Landholders’ Society, was established in 1838 by Dwarkanath Tagore, Prasanna Kumar Tagore, Radhakanta Deb, Ramkamal Sen and Bhabani Charan Mitra.[4][5]
It has been described as “the first organisation of Bengal with distinct political object.” [1] The society virtually became defunct after the death of Dwarkanath Tagore.[4]
British India Society[edit]
The British India Society was set up in 1839 in England primarily as a result of the efforts of William Adam, who had come to India and befriended Ram Mohan Roy. On his return to England he took up India’s cause. Others involved were George Thompson, William Ednis, and Major General Briggs. They organised meetings and enlightened people about conditions in India. In 1841, it started publishing a newspaper named British Indian Advocate, edited by Professor William Adam[1][6]
In 1842, Dwarkanath Tagore went to England, accompanied by Chandramohan Chatterjee and Paramananda Maitra,. It was the second visit abroad by educated Indians and the first since that of Ram Mohan Roy.[6] Dwarkanath Tagore returned from England with George Thompson. His oratory charmed everybody and drew in Young Bengal members such as Ramgopal Ghosh, Tarachand Chakrabarti and Peary Chand Mitra.[7]
As a result of the efforts of George Thompson the Bengal British India Society was founded on 20 April 1843.[8] The object of the society was to secure the welfare, and advance the interests of all classes, but it would “adopt and recommend such measures only as are consistent with pure loyalty to the person and government of the reigning sovereign of the British dominions.” [1]
Its first executive committee consisted of four Europeans and eleven Indians with George Thompson as President, GF Remfry and Ramgopal Ghosh as Vice-Presidents, Peary Chand Mitra as Secretary. The members on the committee were Tarachand Chakrabarti, Dakshinaranjan Mukherjee, Brojonath Dhar, Krishna Mohan Banerjee, Hari Mohan, Govind Chandra Sen, Chandra Sekhar Deb, Shyama Charan Sen and Satkari Datta.[9] There were people like Ramtanu Lahiri, also associated in some way.[10]
In 1849, John Elliot Drinkwater Bethune prepared drafts of certain legislative efforts aimed at bringing the British-born subjects of the crown under the jurisdiction of the British East India Company, but the Europeans named these bills the Black Acts and opposed them tooth and nail.[1][11] This opposition led Indians to think that their fortunes were not linked to those of the Europeans and that they needed associations of Indians to take care of their interests. Moreover, some changes were anticipated with the possible termination of the Company’s charter.[1] Ramgopal Ghosh and Digamber Mitra played a particularly significant role in the formation of the British Indian Society.[4]
Objectives and extent of British Indian Association[edit]
During the early years the activities of the association consisted mainly of submissions of petitions to the Government and to the British Parliament on grievances. There was an inherent trust in the good intentions of the rulers. The association sought to take up issues on behalf of all sections of society but occasionally it made conscious efforts to protect the right of the landed aristocracy.[1] Despite its shortcomings, it was the only association of Indians which took up their causes and represented to the government.[4][12]
From the very beginning, the British Indian Association had an all-India outlook. Associations were formed in different parts of the country and they maintained correspondence with each other. Its first annual report noted with satisfaction the formation at Poona, Madras and Bombay successively of associations of similar nature, “though they elected to carry on operations independently.”[1] In Lucknow, Dakshinaranjan Mukherjee established the Awadh British Indian Association in 1871 and campaigned for the formation of a provincial government with equal number of nominated and elected legislators.[13]
Criticism[edit]
The British Indian Association has been roundly criticised by subsequent political stalwarts. Ambika Charan Mazumdar, a Congress President, wrote: “Constructive policy they had none and seldom, if ever, they laid down any programme of systematic action for the political advancement of the country.”[1]
Bipin Chandra Pal complained that it had failed to cover the country with a network of branches.[1]
However, the British Indian Association played a catalytic role in building up political consciousness in India and more effective organisations followed. The Indian Association was formed in 1876,[1] and the Indian National Congress in 1885.[14]
References[edit]
^ Jump up to: a b c d e f g h i j k l m n Sengupta, Nitish K. (2001). History of the Bengali-speaking People. Ubs Pub Distributors Limited. p. 284. ISBN 978-81-7476-355-6.
Jump up ^ Sastri, Sivanath, Ramtanu Lahiri O Tatkalin Banga Samaj, (in Bengali) 1903/2001, New Age Publishers Pvt. Ltd., p. 116.
Jump up ^ Sastri, Sivanath, pp. 115–6.
^ Jump up to: a b c d Sastri, Sivanath, p. 115.
Jump up ^ Khan, BR (2012). "Zamindari Association, The". In Islam, Sirajul; Jamal, Ahmed A. Banglapedia: National Encyclopedia of Bangladesh (Second ed.). Asiatic Society of Bangladesh.
^ Jump up to: a b Sastri, Sivanath, p. 99.
Jump up ^ Sastri, Sivanath, p. 101.
Jump up ^ Sastri, Sivanath, pp. 101–2.
Jump up ^ Khan, BR (2012). "Bengal British India Society". In Islam, Sirajul; Jamal, Ahmed A. Banglapedia: National Encyclopedia of Bangladesh (Second ed.). Asiatic Society of Bangladesh.
Jump up ^ Sastri, Sivanath, p. 102.
Jump up ^ Sastri, Sivanath, p. 114.
Jump up ^ Mahmood, ABM (2012). "British Indian Association, The". In Islam, Sirajul; Jamal, Ahmed A. Banglapedia: National Encyclopedia of Bangladesh (Second ed.). Asiatic Society of Bangladesh.
Jump up ^ Sengupta, Subodhchandra (1998). Sansad Bangali charitabhidhan. p. 202. ISBN 978-81-85626-65-9.
Jump up ^ Sengupta, Nitish, p. 290.
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Chromosomal crossover
From Wikipedia, the free encyclopedia
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Crossing over occurs between prophase 1 and metaphase 1 and is the process where homologous chromosomes pair up with each other and exchange different segments of their genetic material to form recombinant chromosomes. It can also happen during mitotic division,[1] which may result in loss of heterozygosity. Crossing over is essential for the normal segregation of chromosomes during meiosis.[citation needed] Crossing over also accounts for genetic variation, because due to the swapping of genetic material during crossing over, the chromatids held together by the centromere are no longer identical. So, when the chromosomes go on to meiosis II and separate, some of the daughter cells receive daughter chromosomes with recombined alleles. Due to this genetic recombination, the offspring have a different set of alleles and genes than their parents do. In the diagram, genes B and b are crossed over with each other, making the resulting recombinants after meiosis Ab, AB, ab, and aB.
Thomas Hunt Morgan's illustration of crossing over (1916)
A double crossing over
Chromosomal crossover (or crossing over) is the exchange of genetic material between homologous chromosomes that results in recombinant chromosomes during sexual reproduction. It is one of the final phases of genetic recombination, which occurs in the pachytene stage of prophase I of meiosis during a process called synapsis. Synapsis begins before the synaptonemal complex develops and is not completed until near the end of prophase I. Crossover usually occurs when matching regions on matching chromosomes break and then reconnect to the other chromosome.
Crossing over was described, in theory, by Thomas Hunt Morgan. He relied on the discovery of Frans Alfons Janssens who described the phenomenon in 1909 and had called it "chiasmatypie". The term chiasma is linked, if not identical, to chromosomal crossover. Morgan immediately saw the great importance of Janssens' cytological interpretation of chiasmata to the experimental results of his research on the heredity of Drosophila. The physical basis of crossing over was first demonstrated by Harriet Creighton and Barbara McClintock in 1931.[2]
The linked frequency of crossing over between two gene loci (markers) is the crossing-over value . For fixed set of genetic and environmental conditions, recombination in a particular region of a linkage structure (chromosome) tends to be constant and the same is then true for the crossing-over value which is used in the production of genetic maps.[3][4]
Contents
[hide]
1 Origins
1.1 DNA repair theory
1.2 Links to bacterial transformation
2 Chemistry
2.1 MSH4 and MSH5
2.2 Chiasma
3 Consequences
4 Non-homologous crossover
5 See also
6 References
Origins[edit]
There are two popular and overlapping theories that explain the origins of crossing-over, coming from the different theories on the origin of meiosis. The first theory rests upon the idea that meiosis evolved as another method of DNA repair, and thus crossing-over is a novel way to replace possibly damaged sections of DNA.[5] The second theory comes from the idea that meiosis evolved from bacterial transformation, with the function of propagating diversity.[6]
DNA repair theory[edit]
Crossing over and DNA repair are very similar processes, which utilize many of the same protein complexes.[5][7][8] Recombinases and primases lay a foundation of nucleotides along the DNA sequence. One such particular protein complex that is conserved between processes is RAD51, a well conserved recombinase protein that has been shown to be crucial in DNA repair as well as cross over.[9] Several other genes in D. melanogaster have been linked as well to both processes, by showing that mutants at these specific loci cannot undergo DNA repair or crossing over. Such genes include mei-41, mei-9, hdm, spnA, and brca2.[5] This large group of conserved genes between processes supports the theory of a close evolutionary relationship. Furthermore, DNA repair and crossover have been found to favor similar regions on chromosomes. In an experiment using radiation hybrid mapping on wheat’s (Triticum aestivum L.) 3B chromosome, crossing over and DNA repair were found to occur predominantly in the same regions.[10] Furthermore, crossing over has been correlated to occur in response to stressful, and likely DNA damaging, conditions [11][12]
Links to bacterial transformation[edit]
The process of bacterial transformation also shares many similarities with chromosomal cross over, particularly in the formation of overhangs on the sides of the broken DNA strand, allowing for the annealing of a new strand. Bacterial transformation itself has been linked to DNA repair many times.[5] The second theory comes from the idea that meiosis evolved from bacterial transformation, with the function of propagating genetic diversity.[6] .[13] Thus, this evidence suggests that it is a question of whether cross over is linked to DNA repair or bacterial transformation, as the two do not appear to be mutually exclusive. It is likely that crossing over may have evolved from bacterial transformation, which in turn developed from DNA repair, thus explaining the links between all three processes.
Chemistry[edit]
A current model of meiotic recombination, initiated by a double-strand break or gap, followed by pairing with an homologous chromosome and strand invasion to initiate the recombinational repair process. Repair of the gap can lead to crossover (CO) or non-crossover (NCO) of the flanking regions. CO recombination is thought to occur by the Double Holliday Junction (DHJ) model, illustrated on the right, above. NCO recombinants are thought to occur primarily by the Synthesis Dependent Strand Annealing (SDSA) model, illustrated on the left, above. Most recombination events appear to be the SDSA type.
Meiotic recombination may be initiated by double-stranded breaks that are introduced into the DNA by exposure to DNA damaging agents[5] or the Spo11 protein.[14] One or more exonucleases then digest the 5’ ends generated by the double-stranded breaks to produce 3’ single-stranded DNA tails (see diagram). The meiosis-specific recombinase Dmc1 and the general recombinase Rad51 coat the single-stranded DNA to form nucleoprotein filaments.[15] The recombinases catalyze invasion of the opposite chromatid by the single-stranded DNA from one end of the break. Next, the 3’ end of the invading DNA primes DNA synthesis, causing displacement of the complementary strand, which subsequently anneals to the single-stranded DNA generated from the other end of the initial double-stranded break. The structure that results is a cross-strand exchange, also known as a Holliday junction. The contact between two chromatids that will soon undergo crossing-over is known as a chiasma. The Holliday junction is a tetrahedral structure which can be 'pulled' by other recombinases, moving it along the four-stranded structure.
Holliday Junction
Molecular structure of a Holliday junction.
Molecular structure of a Holliday junction. From PDB: 3CRX.
MSH4 and MSH5[edit]
The MSH4 and MSH5 proteins form a hetero-oligomeric structure (heterodimer) in yeast and humans.[16][17][18] In the yeast Saccharomyces cerevisiae MSH4 and MSH5 act specifically to facilitate crossovers between homologous chromosomes during meiosis.[16] The MSH4/MSH5 complex binds and stabilizes double Holliday junctions and promotes their resolution into crossover products. An MSH4 hypomorphic (partially functional) mutant of S. cerevisiae showed a 30% genome wide reduction in crossover numbers, and a large number of meioses with non exchange chromosomes.[19] Nevertheless, this mutant gave rise to spore viability patterns suggesting that segregation of non-exchange chromosomes occurred efficiently. Thus in S. cerevisiae proper segregation apparently does not entirely depend on crossovers between homologous pairs.
Chiasma[edit]
The grasshopper Melanoplus femur-rubrum was exposed to an acute dose of X-rays during each individual stage of meiosis, and chiasma frequency was measured.[20] Irradiation during the leptotene-zygotene stages of meiosis (that is, prior to the pachytene period in which crossover recombination occurs) was found to increase subsequent chiasma frequency. Similarly, in the grasshopper Chorthippus brunneus, exposure to X-irradiation during the zygotene-early pachytene stages caused a significant increase in mean cell chiasma frequency.[21] Chiasma frequency was scored at the later diplotene-diakinesis stages of meiosis. These results suggest that X-rays induce DNA damages that are repaired by a crossover pathway leading to chiasma formation.
Consequences[edit]
The difference between gene conversion and chromosomal crossover.
In most eukaryotes, a cell carries two versions of each gene, each referred to as an allele. Each parent passes on one allele to each offspring. An individual gamete inherits a complete haploid complement of alleles on chromosomes that are independently selected from each pair of chromatids lined up on the metaphase plate. Without recombination, all alleles for those genes linked together on the same chromosome would be inherited together. Meiotic recombination allows a more independent segregation between the two alleles that occupy the positions of single genes, as recombination shuffles the allele content between homologous chromosomes.
Recombination results in a new arrangement of maternal and paternal alleles on the same chromosome. Although the same genes appear in the same order, some alleles are different. In this way, it is theoretically possible to have any combination of parental alleles in an offspring, and the fact that two alleles appear together in one offspring does not have any influence on the statistical probability that another offspring will have the same combination. This principle of "independent assortment" of genes is fundamental to genetic inheritance.[22] However, the frequency of recombination is actually not the same for all gene combinations. This leads to the notion of "genetic distance", which is a measure of recombination frequency averaged over a (suitably large) sample of pedigrees. Loosely speaking, one may say that this is because recombination is greatly influenced by the proximity of one gene to another. If two genes are located close together on a chromosome, the likelihood that a recombination event will separate these two genes is less than if they were farther apart. Genetic linkage describes the tendency of genes to be inherited together as a result of their location on the same chromosome. Linkage disequilibrium describes a situation in which some combinations of genes or genetic markers occur more or less frequently in a population than would be expected from their distances apart. This concept is applied when searching for a gene that may cause a particular disease. This is done by comparing the occurrence of a specific DNA sequence with the appearance of a disease. When a high correlation between the two is found, it is likely that the appropriate gene sequence is really closer.[23]
Non-homologous crossover[edit]
Crossovers typically occur between homologous regions of matching chromosomes, but similarities in sequence and other factors can result in mismatched alignments. Most DNA is composed of base pair sequences repeated very large numbers of times.[24] These repetitious segments, often referred to as satellites, are fairly homogenous among a species.[24] During DNA replication, each strand of DNA is used as a template for the creation of new strands using a partially-conserved mechanism; proper functioning of this process results in two identical, paired chromosomes, often called sisters. Sister chromatid crossover events are known to occur at a rate of several crossover events per cell per division in eukaryotes. [24] Most of these events involve an exchange of equal amounts of genetic information, but unequal exchanges may occur due to sequence mismatch. These are referred to by a variety of names, including non-homologous crossover, unequal crossover, and unbalanced recombination, and result in an insertion or deletion of genetic information into the chromosome. While rare compared to homologous crossover events, these mutations are drastic, affecting many loci at the same time. They are considered the main driver behind the generation of gene duplications and are a general source of mutation within the genome.[25]
Many proteins are implicated in DSB repair; this figure shows a probable, simplified mechanism.
The specific causes of non-homologous crossover events are unknown, but several influential factors are known to increase the likelihood of an unequal crossover. One common vector leading to unbalanced recombination is the repair of double-strand breaks (DSBs).[26] DSBs are often repaired using non-homologous end joining, a process which involves invasion of a template strand by the DSB strand (see figure below). Nearby homologous regions of the template strand are often used for repair, which can give rise to either insertions or deletions in the genome if a non-homologous but complementary part of the template strand is used.[26] Sequence similarity is a major player in crossover – crossover events are more likely to occur in long regions of close identity on a gene.[27] This means that any section of the genome with long sections of repetitive DNA is prone to crossover events.
The presence of transposable elements is another influential element of non-homologous crossover. Repetitive regions of code characterize transposable elements; complementary but non-homologous regions are ubiquitous within transposons. Because chromosomal regions composed of transposons have large quantities of identical, repetitious code in a condensed space, it is thought that transposon regions undergoing a crossover event are more prone to erroneous complementary match-up;[28] that is to say, a section of a chromosome containing a lot of identical sequences, should it undergo a crossover event, is less certain to match up with a perfectly homologous section of complementary code and more prone to binding with a section of code on a slightly different part of the chromosome. This results in unbalanced recombination, as genetic information may be either inserted or deleted into the new chromosome, depending on where the recombination occurred.
While the motivating factors behind unequal recombination remain obscure, elements of the physical mechanism have been elucidated. Mismatch repair (MMR) proteins, for instance, are a well-known regulatory family of proteins, responsible for regulating mismatched sequences of DNA during replication and escape regulation.[29] The operative goal of MMRs is the restoration of the parental genotype. One class of MMR in particular, MutSβ, is known to initiate the correction of insertion-deletion mismatches of up to 16 nucleotides.[29] Little is known about the excision process in eukaryotes, but E. coli excisions involve the cleaving of a nick on either the 5’ or 3’ strand, after which DNA helicase and DNA polymerase III bind and generate single-stranded proteins, which are digested by exonucleases and attached to the strand by ligase.[29] Multiple MMR pathways have been implicated in the maintenance of complex organism genome stability, and any of many possible malfunctions in the MMR pathway result in DNA editing and correction errors.[30] Therefore, while it is not certain precisely what mechanisms lead to errors of non-homologous crossover, it is extremely likely that the MMR pathway is involved.
See also[edit]
Unequal crossing over
Coefficient of coincidence
Genetic distance
Independent assortment
Mitotic crossover
Recombinant frequency
References[edit]
Jump up ^ Griffiths, AJF; Gelbart, WM; Miller, JH; et al. (1999). "Modern Genetic Analysis: Mitotic Crossing-Over". New York: W. H. Freeman.
Jump up ^ Creighton H, McClintock B (1931). "A Correlation of Cytological and Genetical Crossing-Over in Zea Mays". Proc Natl Acad Sci USA. 17 (8): 492–7. doi:10.1073/pnas.17.8.492. PMC 1076098  . PMID 16587654. (Original paper)
Jump up ^ Rieger R. Michaelis A., Green M. M. (1976). Glossary of genetics and cytogenetics: Classical and molecular. Heidelberg - New York: Springer-Verlag. ISBN 3-540-07668-9. CS1 maint: Uses authors parameter (link)
Jump up ^ King R. C., Stransfield W. D. (1998): Dictionary of genetics. Oxford University Press, New York, Oxford, ISBN 0-19-50944-1-7; ISBN 0-19-509442-5.
^ Jump up to: a b c d e Harris Bernstein, Carol Bernstein and Richard E. Michod (2011). Meiosis as an Evolutionary Adaptation for DNA Repair. Chapter 19 in DNA Repair. Inna Kruman, editor. InTech Open Publisher. doi:10.5772/25117 http://www.intechopen.com/books/dna-repair/meiosis-as-an-evolutionary-adaptation-for-dna-repair
^ Jump up to: a b Bernstein, H; Bernstein, C (2010). "Evolutionary origin of recombination during meiosis". BioScience. 60 (7): 498–505. doi:10.1525/bio.2010.60.7.5.
Jump up ^ Dangel, NJ; Knoll, A; Puchta, H (2014). "MHF1 plays Fanconi anaemia complementation group M protein (FANCM)-dependent and FANCM-independent roles in DNA repair and homologous recombination in plants". Plant J. 78 (5): 822–33. doi:10.1111/tpj.12507. PMID 24635147.
Jump up ^ Saponaro, M; Callahan, D; Zheng, X; Liberi, G (2010). "Cdk1 Targets Srs2 to Complete Synthesis-Dependent Strand Annealing and to Promote Recombinational Repair". PLoS Genet. 6 (2): e1000858. doi:10.1371/journal.pgen.1000858. Retrieved 20 March 2015.
Jump up ^ Esposito, M (September 1978). "Evidence that Spontaneous Mitotic Recombination Occurs at the Two-Strand Stage". Proceedings of the National Academy of Sciences of the USA. 75 (9): 4436–4440. doi:10.1073/pnas.75.9.4436.
Jump up ^ Kumar, A; Bassi, F; Paux, E (2012). "DNA repair and crossing over favor similar chromosome regions as discovered in radiation hybrid of Triticum". BMC Genomics. 13 (339). doi:10.1186/1471-2164-13-339. PMC 3443642  . PMID 22827734. Retrieved 14 March 2015.
Jump up ^ Steinboeck, F (2010). "The relevance of oxidative stress and cytotoxic DNA lesions for spontaneous mutagenesis in non-replicating yeast cells". Mutat Res. 688 (1–2): 47–52. doi:10.1016/j.mrfmmm.2010.03.006. PMID 20223252.
Jump up ^ Nedelcu, M; Marcu, O; Michod, RE (2004). "Sex as a response to oxidative stress: a twofold increase in cellular reactive oxygen species activates sex genes". Proc. R. Soc. B. 271: 1591–1596. doi:10.1098/rspb.2004.2747. PMC 1691771  . PMID 15306305. Retrieved 10 March 2015.
Jump up ^ Charpentier, X (2010). "Antibiotics and UV Radiation Induce Competence for Natural Transformation in Legionella pneumophila". Journal of Bacteriology. 193 (5): 1114–1121. doi:10.1128/JB.01146-10. PMC 3067580  . PMID 21169481. Retrieved 14 March 2015.
Jump up ^ Keeney, S; Giroux, CN; Kleckner, N (1997). "Meiosis-Specific DNA Double-Strand Breaks Are Catalyzed by Spo11, a Member of a Widely Conserved Protein Family". Cell. 88 (3): 375–84. doi:10.1016/S0092-8674(00)81876-0. PMID 9039264.
Jump up ^ Sauvageau, S; Stasiak, Az; Banville, I; Ploquin, M; Stasiak, A; Masson, Jy (June 2005). "Fission Yeast Rad51 and Dmc1, Two Efficient DNA Recombinases Forming Helical Nucleoprotein Filaments" (Free full text). Molecular and Cellular Biology. 25 (11): 4377–87. doi:10.1128/MCB.25.11.4377-4387.2005. ISSN 0270-7306. PMC 1140613  . PMID 15899844.
^ Jump up to: a b Pochart P, Woltering D, Hollingsworth NM (1997). "Conserved properties between functionally distinct MutS homologs in yeast". J. Biol. Chem. 272 (48): 30345–9. doi:10.1074/jbc.272.48.30345. PMID 9374523.
Jump up ^ Winand NJ, Panzer JA, Kolodner RD (1998). "Cloning and characterization of the human and Caenorhabditis elegans homologs of the Saccharomyces cerevisiae MSH5 gene". Genomics. 53 (1): 69–80. doi:10.1006/geno.1998.5447. PMID 9787078.
Jump up ^ Bocker T, Barusevicius A, Snowden T, Rasio D, Guerrette S, Robbins D, Schmidt C, Burczak J, Croce CM, Copeland T, Kovatich AJ, Fishel R (1999). "hMSH5: a human MutS homologue that forms a novel heterodimer with hMSH4 and is expressed during spermatogenesis". Cancer Res. 59 (4): 816–22. PMID 10029069.
Jump up ^ Krishnaprasad GN, Anand MT, Lin G, Tekkedil MM, Steinmetz LM, Nishant KT (2015). "Variation in crossover frequencies perturb crossover assurance without affecting meiotic chromosome segregation in Saccharomyces cerevisiae". Genetics. 199 (2): 399–412. doi:10.1534/genetics.114.172320. PMC 4317650  . PMID 25467183.
Jump up ^ Church K, Wimber DE (1969). "Meiosis in the grasshopper: chiasma frequency after elevated temperature and x-rays". Can. J. Genet. Cytol. 11 (1): 209–16. doi:10.1139/g69-025. PMID 5797806.
Jump up ^ Westerman M (1971). "The effect of x-irradiation on chiasma frequency in Chorthippus brunneus". Heredity. 27 (1): 83–91. doi:10.1038/hdy.1971.73. PMID 5289295.
Jump up ^ "genetic recombination".
Jump up ^ Genetic Recombination
^ Jump up to: a b c Smith, George P. (1976-01-01). "Evolution of Repeated DNA Sequences by Unequal Crossover". Science. 191 (4227): 528–535. doi:10.1126/science.1251186. JSTOR 1741301.
Jump up ^ Graur, Dan; Li, Wen-Hsiung (2000-01-01). Fundamentals of Molecular Evolution. Sinauer. ISBN 9780878932665.
^ Jump up to: a b Puchta, Holger (2005-01-01). "The repair of double-strand breaks in plants: mechanisms and consequences for genome evolution". Journal of Experimental Botany. 56 (409): 1–14. doi:10.1093/jxb/eri025. ISSN 0022-0957. PMID 15557293.
Jump up ^ Metzenberg, Ab; et al. (May 1991). "Homology Requirements for Unequal Crossing Over in Humans". Genetics.
Jump up ^ Robberecht, Caroline; Voet, Thierry; Esteki, Masoud Zamani; Nowakowska, Beata A.; Vermeesch, Joris R. (2012-12-03). "Non-allelic homologous recombination between retrotransposable elements is a driver of de novo unbalanced translocations". Genome Research. 23: gr.145631.112. doi:10.1101/gr.145631.112. ISSN 1088-9051. PMC 3589530  . PMID 23212949.
^ Jump up to: a b c Kunkel, Thomas A.; Erie, Dorothy A. (2005-01-01). "Dna Mismatch Repair". Annual Review of Biochemistry. 74 (1): 681–710. doi:10.1146/annurev.biochem.74.082803.133243. PMID 15952900.
Jump up ^ Surtees, J. A., J.A.; Argueso, J.L.; Alani, E. (2004). "Mismatch Repair Proteins: Key Regulators of Genetic Recombination". Cytogenetic and Genome Research. 107: 146–59. doi:10.1159/000080593.
[hide]
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Genetics: homologous recombination / mobile genetic elements
Primarily prokaryotic
Conjugation
Transduction
Transformation
Occurs in eukaryotes
Transfection
Chromosomal crossover
Gene conversion
Fusion gene
Horizontal gene transfer
Sister chromatid exchange
Transposon
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