Polyploid is a term used to describe cells and organisms containing more than two paired (homologous) sets of chromosomes. Most eukaryotic species are diploid, meaning they have two sets of chromosomes — one set inherited from each parent. However polyploidy is found in some organisms and is especially common in plants. In addition, polyploidy also occurs in some tissues of animals who are otherwise diploid, such as human muscle tissues. This is known as endopolyploidy. (Monoploid organisms also occur; a monoploid has only one set of chromosomes.)

Polyploidy refers to a numerical change in a whole set of chromosomes. Organisms in which a particular chromosome, or chromosome segment, is under- or overrepresented are said to be aneuploid (from the Greek words meaning "not," "good," and "fold"). Therefore the distinction between aneuploidy and polyploidy is that aneuploidy refers to a numerical change in part of the chromosome set, whereas polyploidy refers to a numerical change in the whole set of chromosomes.

Polyploidy may occur due to abnormal cell division, either during mitosis, or commonly during metaphase I in meiosis.

Polyploidy occurs in some animals, such as goldfish, salmon, and salamanders, but is especially common among ferns and flowering plants (see Hibiscus rosa-sinensis), including both wild and cultivated species. Wheat, for example, after millennia of hybridization and modification by humans, has strains that are diploid (two sets of chromosomes), tetraploid (four sets of chromosomes) with the common name of durum or macaroni wheat, and hexaploid (six sets of chromosomes) with the common name of bread wheat. Many agriculturally important plants of the genus Brassica are also tetraploids. Polyploidization is a mechanism of sympatric speciation because polyploids are usually unable to interbreed with their diploid ancestors.

Polyploidy can be induced in plants and cell cultures by some chemicals: the best known is colchicine, which can result in chromosome doubling, though its use may have other less obvious consequences as well. Oryzalin also will double the existing chromosome content.

Polyploid types

Polyploid types are labeled according to the number of chromosome sets in the nucleus: triploid (three sets; 3x), for example seedless watermelons, common in the phylum Tardigrada

tetraploid (four sets; 4x), for example Salmonidae fish

pentaploid (five sets; 5x), for example Kenai Birch (Betula papyrifera var. kenaica)

hexaploid (six sets; 6x), for example wheat, kiwifruit

octaploid (eight sets; 8x), for example Acipenser (genus of sturgeon fish)

decaploid (ten sets; 10x), for example certain strawberries

dodecaploid (twelve sets; 12x), for example the plant Celosia argentea and the amphibian Xenopus ruwenzoriensis

Polyploidy in animals (non-human)

Examples in animals are more common in the 'lower' forms such as flatworms, leeches, and brine shrimp. Polyploid animals are often sterile, so they often reproduce by parthenogenesis. Polyploid lizards are also quite common and parthenogenetic. Polyploid mole salamanders (mostly triploids) are all female and reproduce by kleptogenesis, "stealing" spermatophores from diploid males of related species to trigger egg development but not incorporating the males' DNA into the offspring. While mammalian liver cells are polyploid, rare instances of polyploid mammals are known, but most often result in prenatal death.

One of the few known exceptions to this 'rule' is an octodontid rodent of Argentina's harsh desert regions, known as the Plains Viscacha-Rat (Tympanoctomys barrerae). This rodent is not a rat, but kin to guinea pigs and chinchillas. Its "new" diploid [2n] number is 102 and so its cells are roughly twice normal size. Its closest living relation is Octomys mimax, the Andean Viscacha-Rat of the same family, whose 2n = 56. It is surmised that an Octomys-like ancestor produced tetraploid (i.e., 4n = 112) offspring that were, by virtue of their doubled chromosomes, reproductively isolated from their parents; but that these likely survived the ordinarily catastrophic effects of polyploidy in mammals by shedding (via translocation or some similar mechanism) the "extra" set of sex chromosomes gained at this doubling. (The closely related Golden Vizcacha Rat, 2n = 96, is thought to have arisen via roughly the same process).

Polyploidy in humans

True polyploidy rarely occurs in humans, although it occurs in some tissues (especially in the liver). Aneuploidy is more common.

Polyploidy occurs in humans in the form of triploidy, with 69 chromosomes (sometimes called 69,XXX), and tetraploidy with 92 chromosomes (sometimes called 92,XXXX). Triploidy, usually due to polyspermy, occurs in about 2–3% of all human pregnancies and ~15% of miscarriages. The vast majority of triploid conceptions end as miscarriage and those that do survive to term typically die shortly after birth. In some cases survival past birth may occur longer if there is mixoploidy with both a diploid and a triploid cell population present.

Triploidy may be the result of either digyny (the extra haploid set is from the mother) or diandry (the extra haploid set is from the father). Diandry is mostly caused by reduplication of the paternal haploid set from a single sperm, but may also be the consequence of dispermic (two sperm) fertilization of the egg. Digyny is most commonly caused by either failure of one meiotic division during oogenesis leading to a diploid oocyte or failure to extrude one polar body from the oocyte. Diandry appears to predominate among early miscarriages while digyny predominates among triploidy that survives into the fetal period. However, among early miscarriages, digyny is also more common in those cases <8.5 weeks gestational age or those in which an embryo is present. There are also two distinct phenotypes in triploid placentas and fetuses that are dependent on the origin of the extra haploid set. In digyny there is typically an asymmetric poorly grown fetus, with marked adrenal hypoplasia and a very small placenta. In diandry, a partial hydatidiform mole develops. Huge explosions in angiosperm species diversity appear to have coincided with the timing of ancient genome duplications shared by many species. It has been established that 15% of angiosperm and 31% of fern speciation events are accompanied by ploidy increase. Polyploid plants can arise spontaneously in nature by several mechanisms, including meiotic or mitotic failures, and fusion of unreduced (2n) gametes. Both autopolyploids (e.g. potato) and allopolyploids (e.g. canola, wheat, cotton) can be found among both wild and domesticated plant species. Most polyploids display heterosis relative to their parental species, and may display novel variation or morphologies that may contribute to the processes of speciation and eco-niche exploitation. Many of these rapid changes may contribute to reproductive isolation and speciation.

Lomatia tasmanica is an extremely rare Tasmanian shrub which is triploid and sterile, and reproduction is entirely vegetative with all plants having the same genetic structure.

There are few naturally occurring polyploid conifers. One example is the giant tree Sequoia sempervirens or Coast Redwood which is a hexaploid (6x) with 66 chromosomes (2n = 6x = 66), although the origin is unclear.

Polyploid crops

Polyploid plants tend to be larger and better at flourishing in early succession habitats such as farm fields. In the breeding of crops, the tallest and best thriving plants are selected for. Thus, many crops (and agricultural weeds) may have unintentionally been bred to a higher level of ploidy.

The induction of polyploidy is a common technique to overcome the sterility of a hybrid species during plant breeding. For example, Triticale is the hybrid of wheat (Triticum turgidum) and rye (Secale cereale). It combines sought-after characteristics of the parents, but the initial hybrids are sterile. After polyploidization, the hybrid becomes fertile and can thus be further propagated to become triticale.

In some situations polyploid crops are preferred because they are sterile. For example many seedless fruit varieties are seedless as a result of polyploidy. Such crops are propagated using asexual techniques such as grafting.

Polyploidy in crop plants is most commonly induced by treating seeds with the chemical colchicine.

Examples of polyploid crops

Triploid crops: apple, banana, citrus, ginger, watermelon

Tetraploid crops: apple, durum or macaroni wheat, cotton, potato, cabbage, leek, tobacco, peanut, kinnow, Pelargonium

Hexaploid crops: chrysanthemum, bread wheat, triticale, oat, kiwifruit The preparation and study of karyotypes is part of cytology and, more specifically, cytogenetics.

Although the replication and transcription of DNA is highly standardized in eukaryotes, the same cannot be said for their karotypes, which are highly variable between species in chromosome number and in detailed organization despite being constructed out of the same macromolecules. In some cases there is even significant variation within species. This variation provides the basis for a range of studies in what might be called evolutionary cytology.

Paralogous

The term is used to describe the relationship among duplicated genes or portions of chromosomes that derived from a common ancestral DNA. Paralogous segments of DNA may arise spontaneously by errors during DNA replication, copy and paste transposons, or whole genome duplications.

Homologous

The term is used to describe the relationship of similar chromosomes that pair at mitosis and meiosis. In a diploid, one homolog is derived from the male parent (sperm) and one is derived from the female parent (egg). During meiosis and gametogenesis, homologous chromosomes pair and exchange genetic material by recombination, leading to the production of sperm or eggs with chromosome haplotypes containing novel genetic variation.

Homoeologous

The term homoeologous, also spelled homeologous, is used to describe the relationship of similar chromosomes or parts of chromosomes brought together following inter-species hybridization and allopolyploidization, and whose relationship was completely homologous in an ancestral species. In allopolyploids, the homologous chromosomes within each parental sub-genome should pair faithfully during meiosis, leading to disomic inheritance; however in some allopolyploids, the homoeologous chromosomes of the parental genomes may be nearly as similar to one another as the homologous chromosomes, leading to tetrasomic inheritance (four chromosomes pairing at meiosis), intergenomic recombination, and reduced fertility.

Example of homoeologous chromosomes

Durum wheat is the result of the inter-species hybridization of two diploid grass species Triticum urartu and Aegilops speltoides. Both the diploid ancestors had two sets of 7 chromosomes, which were similar in terms of size and genes contained on them. Durum wheat contains two sets of chromosomes derived from Triticum urartu and two sets of chromosomes derived from Aegilops speltoides. Each chromosome pair derived from the Triticum urartu parent is homoeologous to the opposite chromosome pair derived from the Aegilops speltoides parent, though each chromosome pair unto itself is homologous.

See also

Chromosome

Karyotype

Meiosis

Paleopolyploidy

Polyploid complex

Speciation

Sympatry

References

Further reading

Snustad, P. et al. 2006. Principles of Genetics, 4th ed. John Wiley & Sons, Inc. Hoboken, NJ ISBN 100-471-69939-X

Gregory, T.R. & Mable, B.K. (2005). Polyploidy in animals. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp. 427–517.

Tate, J.A., Soltis, D.E., & Soltis, P.S. (2005). Polyploidy in plants. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp. 371–426.

Van de Peer, Y. and Meyer, A. (2005). Large-scale gene and ancient genome duplications. In The Evolution of the Genome (edited by T.R. Gregory). Elsevier, San Diego, pp. 329–368

External links

Polyploidy on Kimball's Biology Pages

The polyploidy portal a community-editable project with information, research, education, and a bibliography about polyploidy.

Category:Classical genetics Category:Speciation

Coordinates	23°32′34″N46°18′39″N
Consort	yes
Name	Elizabeth Bowes-Lyon
Imgw	200
Caption	The Queen at the 1939 New York World's Fair
Succession	Queen consort of the United Kingdomand the British Dominions;Empress consort of India
Reign	11 December 1936 –6 February 1952
Coronation	12 May 1937
Spouse	George VI
Issue	Elizabeth IIPrincess Margaret, Countess of Snowdon
Full name	Elizabeth Angela Marguerite Bowes-Lyon|group=N}}

| house =House of Windsor (by marriage) | father =Claude Bowes-Lyon, 14th Earl of Strathmore and Kinghorne | mother =Cecilia Cavendish-Bentinck | birth_date =August 04, 1900 | birth_place =London or Hitchin | death_date =March 30, 2002 | death_place =Royal Lodge, Windsor, Berkshire | date of burial =9 April 2002 | place of burial =St. George's Chapel, Windsor Castle }} Elizabeth Bowes-Lyon (4 August 1900 – 30 March 2002) was the Queen consort of King George VI from 1936 until her husband's death in 1952, after which she was known as Queen Elizabeth The Queen Mother, to avoid confusion with her daughter, Queen Elizabeth II. She was the last Queen consort of Ireland and Empress consort of India.

Born into a family of Scottish nobility as The Honourable Elizabeth Bowes-Lyon, she became Lady Elizabeth Bowes-Lyon when her father inherited the Earldom of Strathmore and Kinghorne in 1904. She came to prominence in 1923 when she married Albert, Duke of York, the second son of King George V and Queen Mary. As Duchess of York, she – along with her husband and their two daughters Elizabeth and Margaret – embodied traditional ideas of family and public service. She undertook a variety of public engagements, and became known as the "Smiling Duchess" because of her consistent public expression.

In 1936, her husband unexpectedly became King when her brother-in-law, Edward VIII, abdicated in order to marry the American divorcée Wallis Simpson. As Queen consort, Elizabeth accompanied her husband on diplomatic tours to France and the United States in the run-up to World War II. During the war, her seemingly indomitable spirit provided moral support to the British public. In recognition of her role as an asset to British morale, Adolf Hitler described her as "the most dangerous woman in Europe". After the war, her husband's health deteriorated and she was widowed at the age of 51.

On the death of her mother-in-law Queen Mary in 1953, with her brother-in-law living abroad and her elder daughter Queen at the age of 25, Elizabeth became the senior member of the royal family and assumed a position as family matriarch. In her later years, she was a consistently popular member of the family, when other members were suffering from low levels of public approval. Other possible locations include Forbes House in Ham, London, the home of her maternal grandmother, Mrs Scott. Her birth was registered at Hitchin, Hertfordshire, near the Strathmores' country house, St Paul's Walden Bury, which was also given as her birthplace in the census the following year. She was christened there on 23 September 1900, in the local parish church, All Saints, and her godparents included her paternal aunt Lady Maud Bowes-Lyon and her cousin Mrs Arthur James. In the 1911 census, she was living in Hitchin, but she was not registered as having been born there.

She spent much of her childhood at St Paul's Walden and at Glamis Castle, the Earl's ancestral home in Glamis, Angus, Scotland. She was educated at home by a governess until the age of eight, and was fond of field sports, ponies and dogs. When she started school in London, she astonished her teachers by precociously beginning an essay with two Greek words from Xenophon's Anabasis. Her best subjects were literature and scripture. After returning to private education under a German Jewish governess, Käthe Kübler, she passed the Oxford Local Examination with distinction at age 13.

On her fourteenth birthday, Britain declared war on Germany. Four of her brothers served in the army. Her elder brother, Fergus, an officer in the Black Watch Regiment, was killed in action at the Battle of Loos in 1915. Another brother, Michael, was reported missing in action on 28 April 1917. Three weeks later, the family discovered he had been captured after being wounded. He remained in a prisoner of war camp for the rest of the war. Glamis was turned into a convalescent home for wounded soldiers, which Elizabeth helped to run. She was particularly instrumental in organising the rescue of the Castle's contents during a serious fire on 16 September 1916. One of the soldiers she treated wrote in her autograph book that she was to be "Hung, drawn, & quartered ... Hung in diamonds, drawn in a coach and four, and quartered in the best house in the land."

Marriage to Prince Albert

, 1925.]]

Prince Albert, Duke of York – "Bertie" to the family – was the second son of George V. He initially proposed to Elizabeth in 1921, but she turned him down, being "afraid never, never again to be free to think, speak and act as I feel I really ought to". When he declared he would marry no other, his mother, Queen Mary, visited Glamis to see for herself the girl who had stolen her son's heart. She became convinced that Elizabeth was "the one girl who could make Bertie happy", but nevertheless refused to interfere. At the same time, Elizabeth was courted by James Stuart, Albert's equerry, until he left the prince's service for a better paid job in the American oil business.

In February 1922, Elizabeth was a bridesmaid at the wedding of Albert's sister, Princess Mary, to Viscount Lascelles. The following month, Albert proposed again, but she refused him once more. Eventually, in January 1923, Elizabeth agreed to marry Albert, despite her misgivings about royal life. Albert's freedom in choosing Elizabeth, legally a commoner though the daughter of a peer, was considered a gesture in favour of political modernisation; previously, princes were expected to marry princesses from other royal families. They married on 26 April 1923, at Westminster Abbey. Unexpectedly, Elizabeth laid her bouquet at the Tomb of the Unknown Warrior on her way into the Abbey; a gesture which every royal bride since has copied, though subsequent brides have chosen to do this on the way back from the altar rather than to it. She became styled Her Royal Highness The Duchess of York. Following a wedding breakfast at Buckingham Palace prepared by chef Gabriel Tschumi, they honeymooned at Polesden Lacey, a manor house in Surrey, and then went to Scotland, where she caught "unromantic" whooping cough.

Duchess of York

After a successful visit to Northern Ireland in July 1924, the Labour government agreed that Albert and Elizabeth could tour East Africa from December 1924 to April 1925. The Labour government was defeated by the Conservatives in a general election in November (which Elizabeth described as "marvellous" to her mother) and the Governor-General of Anglo-Egyptian Sudan, Sir Lee Stack, was assassinated three weeks later. Despite this, the tour went ahead, and they visited Aden, Kenya, Uganda, and Sudan, but Egypt was avoided because of political tensions.

Albert had a stammer, which affected his ability to deliver speeches, and after October 1925, Elizabeth assisted in helping him through the therapy devised by Lionel Logue, an episode portrayed in the 2010 film The King's Speech. In 1926, the couple had their first child, Princess Elizabeth – "Lilibet" to the family – who would later become Queen Elizabeth II. Another daughter, Margaret Rose, was born four years later. Albert and Elizabeth, without their child, travelled to Australia to open Parliament House in Canberra in 1927. She was, in her own words, "very miserable at leaving the baby". Their journey by sea took them via Jamaica, the Panama Canal and the Pacific; Elizabeth fretted constantly over her baby back in Britain, but their journey was a public relations success. She charmed the public in Fiji when shaking hands with a long line of official guests, as a stray dog walked in on the ceremony and she shook its paw as well. In New Zealand she fell ill with a cold, and missed some engagements, but enjoyed the local fishing. On the return journey, via Mauritius, the Suez Canal, Malta and Gibraltar, their transport, HMS Renown, caught fire and they prepared to abandon ship before the fire was brought under control.

Accession and abdication of Edward VIII

On 20 January 1936, King George V died and the succession passed to Albert's brother, Prince Edward, the Prince of Wales, who became King Edward VIII. George had expressed reservations about his eldest child, "I pray God that my eldest son will never marry and that nothing will come between Bertie and Lilibet and the throne."

As if granting his father's wish, Edward forced a constitutional crisis by insisting on marrying the American divorcée Mrs Wallis Simpson. Although legally Edward could have married Mrs Simpson, as king he was also head of the Church of England, which at that time did not allow the remarriage of divorced persons. Edward's ministers believed that the people would never accept Mrs. Simpson as queen and advised against the marriage. As a constitutional monarch, Edward was obliged to accept ministerial advice. Rather than abandon his plans to marry Mrs Simpson, Edward chose to abdicate in favour of Albert, who reluctantly became king in his place on 11 December 1936. Albert took the regnal name George VI. He and Elizabeth were crowned King and Queen of Great Britain, Ireland and the British Dominions, and Emperor and Empress of India on 12 May 1937, the date already nominated for the coronation of Edward VIII. Elizabeth's crown was made of platinum and contained the Koh-i-Noor diamond. Edward and Mrs Simpson married and became the Duke and Duchess of Windsor, but while Edward was a Royal Highness, George VI decided to withhold the style from the Duchess, a decision which Elizabeth supported. Elizabeth was later quoted as referring to the Duchess as "that woman".

Queen consort

State visits and royal tour

in New York City]] In summer 1938, a state visit to France by the King and Queen was postponed for three weeks because of the death of the Queen's mother, Lady Strathmore. In two weeks, Norman Hartnell created an all white trousseau for the Queen, who could not wear colours as she was still in mourning. The visit was designed to bolster Anglo-French solidarity in the face of aggression from Nazi Germany. The French press praised the demeanour and charm of the royal couple during the delayed but successful visit, augmented by Hartnell's wardrobe.

Nevertheless, Nazi aggression continued, and the government prepared for war. After the Munich Agreement of 1938 appeared to forestall the advent of armed conflict, the British Prime Minister Neville Chamberlain was invited onto the balcony of Buckingham Palace with the King and Queen to receive acclamation from a crowd of well-wishers. While broadly popular among the general public, Chamberlain's policy towards Hitler was the subject of some opposition in the House of Commons, which led historian John Grigg to describe the King's behaviour in associating himself so prominently with a politician as "the most unconstitutional act by a British sovereign in the present century". However, historians have also argued that the King only ever followed ministerial advice and acted as he was constitutionally bound to do.

In June 1939, Elizabeth and her husband toured North America. The tour was designed to bolster trans-Atlantic support in the event of war, and to affirm Canada's status as a self-governing kingdom sharing with Britain the same person as monarch. The tour took them across Canada from coast to coast and back, and into the United States, where they visited the Roosevelts in the White House and at their Hudson Valley estate. According to an often-told story, during one of the earliest of the royal couple's repeated encounters with the crowds, a Second Boer War veteran asked Elizabeth, "Are you Scots or are you English?" She replied, "I am a Canadian!" Their reception by the Canadian and U.S. public was extremely enthusiastic, and largely dissipated any residual feeling that George and Elizabeth were a lesser substitute for Edward. More critically, U.S. First Lady Eleanor Roosevelt said that Elizabeth was "perfect as a Queen, gracious, informed, saying the right thing & kind but a little self-consciously regal". Elizabeth told Prime Minister Mackenzie King, "that tour made us", and she returned to Canada frequently both on official tours and privately.

World War II

.]] During World War II, the King and Queen became symbols of the nation's determination to fight fascism. Shortly after the declaration of war, The Queen's Book of the Red Cross was conceived. Fifty authors and artists contributed to the book, which was fronted by Cecil Beaton's portrait of the Queen and was sold in aid of the Red Cross. Elizabeth publicly refused to leave London or send the children to Canada, even during the Blitz, when she was advised by the Cabinet to do so. She said, "The children won't go without me. I won't leave the King. And the King will never leave."

She visited troops, hospitals, factories, and parts of Britain that were targeted by the German Luftwaffe, in particular the East End, near London's docks. Her visits initially provoked hostility. Rubbish was thrown at her and the crowds jeered, in part because she dressed in expensive clothing which served to alienate her from those suffering the privations caused by the war. She explained that if the public came to see her they would wear their best clothes, so she should reciprocate in kind; Norman Hartnell dressed her in gentle colours and never black, in order to represent "the rainbow of hope". When Buckingham Palace itself took several hits during the height of the bombing, Elizabeth was able to say, "I'm glad we've been bombed. It makes me feel I can look the East End in the face."

Though the King and Queen spent the working day at Buckingham Palace, partly for security and family reasons they stayed at night at Windsor Castle about west of central London with the Princesses Elizabeth and Margaret. The Palace had lost much of its staff to the army, and most of the rooms were shut. The windows were shattered by bomb blasts, and had to be boarded up. During the "Phoney War" the Queen was given revolver training because of fears of imminent invasion.

Because of her effect on British morale, Adolf Hitler is said to have called her "the most dangerous woman in Europe". At the end of the war in 1945, Churchill was invited onto the balcony in a similar gesture to that given to Chamberlain.

Post-war years

In the 1945 British general election, Churchill's Conservative party was soundly defeated by the Labour party of Clement Attlee. Elizabeth's political views were rarely disclosed, but a letter she wrote in 1947 described Attlee's "high hopes of a socialist heaven on earth" as fading and presumably describes those who voted for him as "poor people, so many half-educated and bemused. I do love them." Woodrow Wyatt thought her "much more pro Conservative" than other members of the royal family, but she later told him, "I like the dear old Labour Party." She also told the Duchess of Grafton, "I love communists". After six years in office, Attlee was defeated in the 1951 British general election and Churchill returned to power.

During the 1947 royal tour of South Africa, Elizabeth's serene public behaviour was broken, exceptionally, when she rose from the royal car to strike an admirer with her umbrella because she had mistaken his enthusiasm for hostility. The 1948 royal tour of Australia and New Zealand was postponed because the King was suffering from increasing ill health. In March 1949, he had a successful operation to improve the circulation in his right leg. In summer 1951, Queen Elizabeth and her daughters fulfilled the King's public engagements in his place. In September, he was diagnosed with lung cancer. After a lung resection, he appeared to recover, but the delayed trip to Australia and New Zealand was altered so that Princess Elizabeth and her husband, the Duke of Edinburgh, went in the King and Queen's place.

Queen Mother

Widowhood

, by Allan Warren.]]

On 6 February 1952, King George VI died peacefully in his sleep. Shortly afterward, Elizabeth began to be styled Her Majesty Queen Elizabeth The Queen Mother. This style was adopted because the normal style for the widow of a king, "Queen Elizabeth", would have been too similar to the style of her elder daughter, now Queen Elizabeth II. Popularly, she simply became the "Queen Mother" or the "Queen Mum".

She was devastated by the King's death and retired to Scotland. However, after a meeting with the Prime Minister, Winston Churchill, she broke her retirement and resumed her public duties. Eventually she became just as busy as Queen Mother as she had been as Queen. In July 1953, she undertook her first overseas visit since the funeral when she laid the foundation stone of the University College of Rhodesia and Nyasaland – the current University of Zimbabwe in Mount Pleasant. She returned in 1957 when she was inaugurated as the College's President, and attended other events in the region that were deliberately designed to be multi-racial. During her daughter's extensive tour of the Commonwealth over 1953–54, Elizabeth acted as a Counsellor of State and looked after her grandchildren, Charles and Anne.

The widowed queen oversaw the restoration of the remote Castle of Mey on the Caithness coast of Scotland, which she used to "get away from everything" for three weeks in August and ten days in October each year. Inspired by the amateur jockey Lord Mildmay, she developed an interest in horse racing, particularly steeplechasing, that continued for the rest of her life. She owned the winners of approximately 500 races. Her distinctive colours of blue with buff stripes were carried by horses such as Special Cargo, the winner of the 1984 Whitbread Gold Cup, and Devon Loch, which spectacularly halted just short of the winning post at the 1956 Grand National. Although (contrary to rumour) she never placed bets, she did have the racing commentaries piped direct to her London residence, Clarence House, so she could follow the races. As an art collector, she purchased works by Claude Monet, Augustus John and Peter Carl Fabergé, among others.

In February 1964, she had an emergency appendectomy, which led to the postponement of a planned tour of Australia, New Zealand and Fiji until 1966. She recuperated during a Caribbean cruise aboard the royal yacht, Britannia. In December 1966, she underwent an operation to remove a tumour after she was diagnosed with colon cancer. Contrary to rumours, she did not have a colostomy. In 1982, she was rushed to hospital when a fish bone stuck in her throat, and had an operation to remove it. Being a keen angler, she calmly joked afterwards, "The salmon have got their own back." In 1984, she had a second operation for cancer, when a lump was removed from her breast, and a second gastric obstruction in 1986 cleared without the need for an operation, but she was hospitalised overnight.

In 1975, she visited Iran at the invitation of Shah Reza Pahlavi. The British ambassador and his wife, Anthony and Sheila Parsons, noted how the Iranians were bemused by her habit of speaking to everyone regardless of status or importance, and hoped the Shah's entourage would learn from the visit to pay more attention to ordinary people. Four years later, the Shah was deposed. Between 1976 and 1984, she made annual summer visits to France, which were among 22 private trips to continental Europe between 1963 and 1992.

Before the marriage of Lady Diana Spencer to her grandson Prince Charles, and after Diana's death, Queen Elizabeth – known for her personal and public charm – was by far the most popular member of the royal family. In 1995, she attended events commemorating the end of the war fifty years before, and had two operations: one to remove a cataract in her left eye, and one to replace her right hip. In 1998, her left hip was replaced after it was broken when she slipped and fell during a visit to Sandringham stables. Her 100th birthday was celebrated in a number of ways: a parade that celebrated the highlights of her life included contributions from Norman Wisdom and John Mills; her image appeared on a special commemorative £20 note issued by the Royal Bank of Scotland; and she attended a lunch at the Guildhall, London, at which George Carey, the Archbishop of Canterbury, accidentally attempted to drink her glass of wine. Her quick admonition of "That's mine!" caused widespread amusement. In November 2000, she broke her collar bone in a fall that kept her recuperating at home over Christmas and the New Year.

In December 2001 aged 101, the Queen Mother had a fall in which she fractured her pelvis. Even so, she insisted on standing for the National Anthem during the memorial service for her husband on 6 February the following year. Just three days later, her second daughter Princess Margaret died. On 13 February 2002, the Queen Mother fell and cut her arm at Sandringham House. Despite this fall, the Queen Mother was still determined to attend Margaret's funeral at St. George's Chapel, Windsor Castle, two days later on Friday of that week. The Queen and the rest of the royal family were greatly concerned about the journey the Queen Mother was facing to get from Norfolk to Windsor. Nevertheless, she made the journey but insisted that she be shielded from the press, so that no photographs of her in a wheelchair could be taken. More than 200,000 people over three days filed past as she lay in state in Westminster Hall at the Palace of Westminster. Members of the household cavalry and other branches of the armed forces stood guard at the four corners of the catafalque. At one point, the Queen Mother's four grandsons Prince Charles, Prince Andrew, Prince Edward and Viscount Linley mounted the guard as a mark of respect known as the Vigil of the Princes—a very high honour only bestowed once before, at King George V's lying in state.

On the day of the Queen Mother's funeral, 9 April, the Governor General of Canada issued a proclamation asking Canadians to honour on that day the memory of the late queen consort. In Australia, the Governor-General read the lesson at the memorial service for the Queen Mother, held in St. Andrew's Cathedral, Sydney. In London, more than a million people filled the area outside Westminster Abbey and along the route from central London to her final resting place beside her husband and younger daughter in St. George's Chapel, Windsor Castle. At her request, after her funeral the wreath that had lain atop her coffin was placed on the Tomb of the Unknown Warrior in Westminster Abbey, a gesture that echoed her wedding-day tribute.

Public perception

Despite being regarded as one of the most popular members of the royal family in recent times who helped to stabilise the popularity of the monarchy as a whole, Elizabeth was subject to various degrees of criticism during her life.

Allegations that during World War II Elizabeth did not abide by the rationing regulations to which the rest of the population was subject, are contradicted by the official records; Eleanor Roosevelt during her stay at Buckingham Palace during the war reported expressly on the rationed food served in the Palace and the limited bathwater that was permitted.

Further allegations that Elizabeth used racist slurs to refer to black people Major Burgess was the husband of Elizabeth Burgess, a mixed-race secretary who accused members of the Prince of Wales's Household of racial abuse. Queen Elizabeth made no public comments on race, but according to Robert Rhodes James in private she "abhorred racial discrimination" and decried apartheid as "dreadful". Woodrow Wyatt records in his diary that when he expressed the view that non-white countries have nothing in common with "us", she told him, "I am very keen on the Commonwealth. They're all like us." However, she did distrust Germans; she told Woodrow Wyatt, "Never trust them, never trust them." While she may have held such views, it has been argued that they were normal for British people of her generation and upbringing, who had experienced two vicious wars with Germany.

In 1987, she was criticised when it emerged that two of her nieces, Katherine Bowes-Lyon and Nerissa Bowes-Lyon, had both been committed to a psychiatric hospital because they were severely handicapped. However, Burke's Peerage had listed the sisters as dead, apparently because their mother, Fenella (the Queen Mother's sister-in-law), "was 'extremely vague' when it came to filling in forms and might not have completed the paperwork for the family entry correctly". When Nerissa had died the year before, her grave was originally marked with a plastic tag and a serial number. The Queen Mother claimed that the news of their institutionalisation came as a surprise to her.

Legacy

Sir Hugh Casson said she was like "a wave breaking on a rock, because although she is sweet and pretty and charming, she also has a basic streak of toughness and tenacity. ... when a wave breaks on a rock, it showers and sparkles with a brilliant play of foam and droplets in the sun, yet beneath is really hard, tough rock, fused, in her case, from strong principles, physical courage and a sense of duty." Peter Ustinov described her during a student demonstration at the University of Dundee in 1968, "As we arrived in a solemn procession the students pelted us with toilet rolls. They kept hold of one end, like streamers at a ball, and threw the other end. The Queen Mother stopped and picked these up as though somebody had misplaced them. [Returning them to the students she said,] 'Was this yours? Oh, could you take it?' And it was her sang-froid and her absolute refusal to be shocked by this, which immediately silenced all the students. She knows instinctively what to do on those occasions. She doesn't rise to being heckled at all; she just pretends it must be an oversight on the part of the people doing it. The way she reacted not only showed her presence of mind, but was so charming and so disarming, even to the most rabid element, that she brought peace to troubled waters."

She was well-known for her dry witticisms. On hearing that Edwina Mountbatten was buried at sea, she said: "Dear Edwina, she always liked to make a splash." After being advised by a Conservative Minister in the 1970s not to employ homosexuals, the Queen Mother observed that without them, "we'd have to go self-service". Emine Saner of The Guardian suggests that with a gin and Dubonnet at noon, red wine with lunch, a port and martini at 6 pm and two glasses of champagne at dinner, "a conservative estimate puts the number of alcohol units she drank at 70 a week". Her extravagant lifestyle amused journalists, particularly when it was revealed she had a multi-million pound overdraft with Coutts Bank.

Her habits were often parodied (with relative affection) by the satirical 1980s television programme Spitting Image – which portrayed her with a Birmingham accent (modelled on actress Beryl Reid) and an ever-present copy of the Racing Post. She was portrayed in the 2002 television film Bertie and Elizabeth by Juliet Aubrey, the 2006 film The Queen by Sylvia Syms and in the 2010 film The King's Speech by Helena Bonham Carter, who was nominated for an Academy Award for Best Supporting Actress and won a BAFTA Award for Best Actress in a Supporting Role for her portrayal as the Queen Mother. , near Toronto, with the effigies of Queen Elizabeth and King George VI.]] The Queen Mother left her entire estate to the Queen, except for some bequests to members of her staff. Her estate was estimated to be worth £70 million, including paintings, Fabergé eggs, jewellery, and horses. Eight years before her death, she had reportedly placed two-thirds of her money into trusts, for the benefit of her great-grandchildren. The Queen Mother's most important pieces of art were transferred to the Royal Collection by the Queen.

A statue of Queen Elizabeth by sculptor Philip Jackson at the George VI Memorial, off The Mall, London, was unveiled on 24 February 2009. The Cunard White Star Line's RMS Queen Elizabeth was named after Elizabeth. She launched the ship on 27 September 1938 in Clydebank, Scotland. Supposedly, the liner started to slide into the water before Elizabeth could officially launch her, and acting sharply, the Queen managed to smash a bottle of Australian red wine over the liner's bow just before she slid out of reach. In 1954, Elizabeth sailed to New York on her namesake.

In March 2011 her eclectic musical taste was revealed when details of her small record collection kept at at the Castle of Mey were made public. She had a taste for ska music and her records included artists such as the yodelling Montana Slim, Tony Hancock, The Goons and Noël Coward. Other music included local folk, Scottish reels and the musicals Oklahoma! and The King and I.

Titles, styles, honours and arms

Titles and styles

Elizabeth held a number of different titles and styles throughout her life, as the daughter of an earl, through her husband, and eventually as consort to the sovereign of multiple states. As consort, she was commonly The Queen. In conversation, the practice was to initially address her as Your Majesty and thereafter as Ma'am.

Arms

The Queen Mother's coat of arms were the Royal Coat of Arms of the United Kingdom (in either the English or the Scottish version) impaled with the arms of her father, the Earl of Strathmore; the latter being 1st and 4th quarters, argent, a lion rampant Azure, armed and langued gules, within a double tressure flory-counter-flory of the second (Lyon), 2nd and 3rd quarters, ermine three bows, stringed paleways proper (Bowes). The shield is surmounted by the imperial crown, and supported by the crowned lion of England and a "lion per fess Or and Gules".

Ancestry

Notes

References

Bibliography

Goldman, Lawrence (May 2006) "Elizabeth (1900–2002)", Oxford Dictionary of National Biography, Oxford University Press, , retrieved 1 May 2009 (Subscription required)

External links

Official site

Remember This—An Elegy on the death of HM Queen Elizabeth, The Queen Mother by Andrew Motion, Poet Laureate, at the BBC

Order of Canada Citation

CBC Digital Archives – Their Majesties in Canada: The 1939 Royal Tour

The Queen Mother's Wedding on The Royal Channel on YouTube.com

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