title: Djuna Barnes

text: In 1912 Barnes's family, facing financial ruin, split up. Elizabeth moved to New York City with Barnes and three of her brothers, then filed for divorce, freeing Wald to marry Fanny Clark. The move gave Barnes an opportunity to study art formally for the first time; she attended the Pratt Institute for about six months from 1912 to 1913 and at the Art Student's League of New York from 1915 to 1916, but the need to support herself and her family—a burden that fell largely on her—soon drove her to leave school and take a job as a reporter at the "Brooklyn Daily Eagle". Upon arriving at the "Daily Eagle", Barnes declared, "I can draw and write, and you'd be a fool not to hire me", words that were inscribed inside the Brooklyn Museum.


title: Djuna Barnes

text: Over the next few years her work appeared in almost every newspaper in New York, including the "New York Press", "The World" and "McCall's"; she wrote interviews, features, theatre reviews, and a variety of news stories, often illustrating them with her own drawings. She also published short fiction in the "New York Morning Telegraph"s Sunday supplement and in the pulp magazine "All-Story Cavalier Weekly".


title: Djuna Barnes

text: Much of Barnes's journalism was subjective and experiential. Writing about a conversation with James Joyce, she admitted to missing part of what he said because her attention had wandered, though she revered Joyce's writing. Interviewing the successful playwright Donald Ogden Stewart, she shouted at him for "roll[ing] over and find[ing] yourself famous" while other writers continued to struggle, then said she wouldn't mind dying; as her biographer Phillip Herring points out, this is "a depressing and perhaps unprecedented note on which to end an interview." For "The Girl and the Gorilla", published by "New York World Magazine" in October 1914 she has a conversation with Dinah, a female gorilla at the Bronx Zoo.


title: Djuna Barnes

text: For another article in "New York World" in 1914, she submitted to force-feeding, a technique then being used on hunger-striking suffragists. Barnes wrote "If I, play acting, felt my being burning with revolt at this brutal usurpation of my own functions, how they who actually suffered the ordeal in its acutest horror must have flamed at the violation of the sanctuaries of their spirits." She concluded "I had shared the greatest experience of the bravest of my sex."


title: Djuna Barnes

text: While she mocked conservative suffrage activist Carrie Chapman Catt when Catt admonished would-be suffrage orators never to "hold a militant pose", or wear "a dress that shows your feet in front", Barnes was supportive of progressive suffragists. Barnes suggested that Catt's conservatism was an obstacle to the suffrage movement when Catt tried to ostracize fellow suffragists Alice Paul and Lucy Burns, who sought the vote for women through media attention directed at their strikes and non-violent protesting. It was their mistreatment that motivated Barnes to experience for herself the torture of being force-fed.


title: Djuna Barnes

text: Barnes immersed herself in risky situations in order to access experiences that a previous generation of homebound women had been denied. Writing about the traditionally masculine domain of boxing from the ringside, Barnes explored boxing as a window into women's modern identities. In 1914, she first posed the question "What do women want at a fight?" in an article titled "My Sisters and I at a New York Prizefight" published in "New York World" magazine. According to Irene Gammel, "Barnes' essay effectively begins to unravel an entire cultural history of repression for women". Barnes's interest in boxing continued into 1915 when she interviewed heavyweight champion Jess Willard.


title: Djuna Barnes

text: In 1915 Barnes moved out of her family's flat to an apartment in Greenwich Village, where she entered a thriving Bohemian community of artists and writers. Among her social circle were Edmund Wilson, Berenice Abbott, and the Dadaist artist and poet Elsa von Freytag-Loringhoven, whose biography Barnes tried to write but never finished. She came into contact with Guido Bruno, an entrepreneur and promoter who published magazines and chapbooks from his garret on Washington Square. Bruno had a reputation for unscrupulousness, and was often accused of exploiting Greenwich Village residents for profit—he used to charge tourists admission to watch Bohemians paint—but he was a strong opponent of censorship and was willing to risk prosecution by publishing Barnes's 1915 collection of "rhythms and drawings"


title: Djuna Barnes

text: Despite a description of sex between women in the first poem, the book was never legally challenged; the passage seems explicit now, but at a time when lesbianism was virtually invisible in American culture, the New York Society for the Suppression of Vice may not have understood its imagery. Others were not as naïve, and Bruno was able to cash in on the book's reputation by raising the price from fifteen to fifty cents and pocketing the difference. Twenty years later Barnes used Bruno as one of the models for Felix Volkbein in "Nightwood", caricaturing his pretensions to nobility and his habit of bowing down before anyone titled or important.


title: Djuna Barnes

text: Barnes was a member of the Provincetown Players, an amateur theatrical collective whose emphasis on artistic rather than commercial success meshed well with her own values. The Players' Greenwich Village theater was a converted stable with bench seating and a tiny stage; according to Barnes it was "always just about to be given back to the horses." Yet it played a significant role in the development of American drama, featuring works by Susan Glaspell, Edna St. Vincent Millay, Wallace Stevens, and Theodore Dreiser, as well as launching the career of Eugene O'Neill. Three one-act plays by Barnes were produced there in 1919 and 1920; a fourth, "The Dove", premiered at Smith College in 1925, and a series of short closet dramas were published in magazines, some under Barnes's pseudonym Lydia Steptoe.


title: Djuna Barnes

text: These plays show the strong influence of the Irish playwright J. M. Synge; she was drawn to both the poetic quality of Synge's language and the pessimism of his vision. Critics have found them derivative, particularly those in which she tried to imitate Synge's Irish dialect, and Barnes may have agreed, since in later years she dismissed them as mere juvenilia. Yet in their content, these stylized and enigmatic early plays are more experimental than those of her fellow playwrights at Provincetown. A "New York Times" review by Alexander Woollcott of her play "Three From the Earth" called it a demonstration of "how absorbing and essentially dramatic a play can be without the audience ever knowing what, if anything, the author is driving at ... The spectators sit with bated breath listening to each word of a playlet of which the darkly suggested clues leave the mystery unsolved."


title: Djuna Barnes

text: Greenwich Village in the 1910s was known for its atmosphere of sexual as well as intellectual freedom. Barnes was unusual among Villagers in having been raised with a philosophy of free love, espoused both by her grandmother and her father. Her father's idiosyncratic vision had included a commitment to unlimited procreation, which she strongly rejected; criticism of childbearing would become a major theme in her work. She did, however, retain sexual freedom as a value. In the 1930s she told Antonia White that "she had no feeling of guilt whatever about sex, about going to bed with any man or woman she wanted"; correspondence indicates that by the time she was 21 her family was well aware of her bisexuality, and she had a number of affairs with both men and women during her Greenwich Village years.


title: Djuna Barnes

text: Of these, the most important was probably her engagement to Ernst Hanfstaengl, a Harvard graduate who ran the American branch of his family's art publishing house. Hanfstaengl had once given a piano concert at the White House and was a friend of then-New York State Senator Franklin Delano Roosevelt, but he became increasingly angered by anti-German sentiment in the United States during World War I. In 1916, he told Barnes he wanted a German wife; the painful breakup became the basis of a deleted scene in "Nightwood". He later returned to Germany and became a close associate of Adolf Hitler. Starting in 1916 or 1917, she lived with a socialist philosopher and critic named Courtenay Lemon, whom she referred to as her common-law husband, but this too ended, for reasons that are unclear. She also had a passionate romantic relationship with Mary Pyne, a reporter for the "New York Press" and fellow member of the Provincetown Players. Pyne died of tuberculosis in 1919, attended by Barnes until the end.


title: Djuna Barnes

text: In the 1920s, Paris was the center of modernism in art and literature. Barnes first traveled there in 1921 on an assignment for "McCall's". She interviewed her fellow expatriate writers and artists for U.S. periodicals and soon became a well-known figure on the local scene; her black cloak and her acerbic wit are remembered in many memoirs of the time. Even before her first novel was published, her literary reputation was already high, largely on the strength of her story "A Night Among the Horses," which was published in "The Little Review" and reprinted in her 1923 collection "A Book". She was part of the inner circle of the influential salon hostess Natalie Barney, who became a lifelong friend and patron, as well as the central figure in Barnes's satiric chronicle of Paris lesbian life, "Ladies Almanack." The most important relationship of Barnes's Paris years was with the artist Thelma Wood. Wood was a Kansas native who had come to Paris to become a sculptor, but at Barnes's suggestion took up silverpoint instead, producing drawings of animals and plants that one critic compared to Henri Rousseau. By the winter of 1922 they had set up housekeeping together in a flat on the Boulevard Saint-Germain. Another close friendship that developed during this time was with the Dada artist Baroness Elsa von Freytag-Loringhoven, with whom Barnes began an intensive correspondence in 1923. "Where Wood gave Barnes a doll as a gift to represent their symbolic love child, the Baroness proposed an erotic marriage whose love-child would be their book." From Paris, Barnes supported the Baroness in Berlin with money, clothing, and magazines. She also collected the Baroness's poems and letters.


title: Djuna Barnes

text: Barnes arrived in Paris with a letter of introduction to James Joyce, whom she interviewed for "Vanity Fair" and who became a friend. The headline of her "Vanity Fair" interview billed him as "the man who is, at present, one of the more significant figures in literature," but her personal reaction to "Ulysses" was less guarded: "I shall never write another line ... Who has the nerve to after that?" It may have been reading Joyce that led Barnes to turn away from the late 19th century Decadent and Aesthetic influences of "The Book of Repulsive Women" toward the modernist experimentation of her later work. They differed, however, on the proper subject of literature; Joyce thought writers should focus on commonplace subjects and make them extraordinary, while Barnes was always drawn to the unusual, even the grotesque. Then, too, her own life was an extraordinary subject. Her autobiographical first novel "Ryder" would not only present readers with the difficulty of deciphering its shifting literary styles—a technique inspired by "Ulysses"—but also with the challenge of piecing together the history of an unconventional polygamous household, far removed from most readers' expectations and experience.


title: Djuna Barnes

text: Despite the difficulties of the text, "Ryder"s bawdiness drew attention, and it briefly became a "New York Times" bestseller. Its popularity caught the publisher unprepared; a first edition of 3,000 sold out quickly, but by the time more copies made it into bookstores, public interest in the book had died down. Still, the advance allowed Barnes to buy a new apartment on Rue Saint-Romain, where she lived with Thelma Wood starting in September 1927. The move made them neighbors of Mina Loy, a friend of Barnes's since Greenwich Village days, who appeared in "Ladies Almanack" as Patience Scalpel, the sole heterosexual character, who "could not understand Women and their Ways."


title: Djuna Barnes

text: Due to its subject matter, "Ladies Almanack" was published in a small, privately printed edition under the pseudonym "A Lady of Fashion." Copies were sold on the streets of Paris by Barnes and her friends, and Barnes managed to smuggle a few into the United States to sell. A bookseller, Edward Titus, offered to carry "Ladies Almanack" in his store in exchange for being mentioned on the title page, but when he demanded a share of the royalties on the entire print run, Barnes was furious. She later gave the name Titus to the abusive father in "The Antiphon".


title: Djuna Barnes

text: Barnes dedicated "Ryder" and "Ladies Almanack" to Thelma Wood, but the year both books were published—1928—was also the year that she and Wood separated. Barnes had wanted their relationship to be monogamous, but had discovered that Wood wanted her "along with the rest of the world." Wood had a worsening dependency on alcohol, and she spent her nights drinking and seeking out casual sex partners; Barnes would search the cafés for her, often winding up equally drunk. Barnes broke up with Wood over her involvement with heiress Henriette McCrea Metcalf (1888–1981), who would be scathingly portrayed in "Nightwood" as Jenny Petherbridge.


title: Djuna Barnes

text: Much of "Nightwood" was written during the summers of 1932 and 1933, while Barnes was staying at Hayford Hall, a country manor in Devon rented by the art patron Peggy Guggenheim. Fellow guests included Antonia White, John Ferrar Holms, and the novelist and poet Emily Coleman. Evenings at the manor—nicknamed "Hangover Hall" by its residents—often featured a party game called Truth that encouraged brutal frankness, creating a tense emotional atmosphere. Barnes was afraid to leave her work in progress unattended because the volatile Coleman, having told Barnes one of her secrets, had threatened to burn the manuscript if Barnes revealed it. But once she had read the book, Coleman became its champion. Her critiques of successive drafts led Barnes to make major structural changes, and when publisher after publisher rejected the manuscript, it was Coleman who pressed T. S. Eliot, then an editor at Faber and Faber, to read it.


title: Djuna Barnes

text: Faber published the book in 1936. Though reviews treated it as a major work of art, the book did not sell well. Barnes received no advance from Faber and the first royalty statement was for only £43; the U.S. edition published by Harcourt, Brace the following year fared no better. Barnes had published little journalism in the 1930s and was largely dependent on Peggy Guggenheim's financial support. She was constantly ill and drank more and more heavily—according to Guggenheim, she accounted for a bottle of whiskey per day. In February 1939 she checked into a hotel in London and attempted suicide. Guggenheim funded hospital visits and doctors, but finally lost patience and sent her back to New York. There she shared a single room with her mother, who coughed all night and who kept reading her passages from Mary Baker Eddy, having converted to Christian Science. In March 1940, her family sent her to a sanatorium in upstate New York to dry out. Furious, Barnes began to plan a biography of her family, writing to Emily Coleman that "there is no reason any longer why I should feel for them in any way but hate." This idea would eventually come to fruition in her play "The Antiphon". After she returned to New York City, she quarreled bitterly with her mother and was thrown out on the street.


title: Djuna Barnes

text: Left with nowhere else to go, Barnes stayed at Thelma Wood's apartment while Wood was out of town, then spent two months on a working ranch in Arizona with Emily Coleman and Coleman's lover Jake Scarborough. She returned to New York and, in September, moved into the small apartment at 5 Patchin Place in Greenwich Village where she would spend the last 41 years of her life. Throughout the 1940s, she continued to drink and wrote virtually nothing. Guggenheim, despite misgivings, provided her with a small stipend, and Coleman, who could ill afford it, sent US$20 per month (about $310 in 2011). In 1943, Barnes was included in Peggy Guggenheim's show "Exhibition by 31 Women" at the Art of This Century gallery in New York.


title: Djuna Barnes

text: In 1946, she worked for Henry Holt as a manuscript reader, but her reports were invariably caustic, and she was fired.


title: Djuna Barnes

text: In 1950, realizing that alcoholism had made it impossible for her to function as an artist, Barnes stopped drinking in order to begin work on her verse play "The Antiphon". The play drew heavily on her own family history, and the writing was fueled by anger; she said "I wrote "The Antiphon" with clenched teeth, and I noted that my handwriting was as savage as a dagger." When he read the play, her brother Thurn accused her of wanting "revenge for something long dead and to be forgotten". Barnes, in the margin of his letter, described her motive as "justice", and next to "dead" she inscribed, "not dead".


title: Djuna Barnes

text: After "The Antiphon", Barnes returned to writing poetry, which she worked and reworked, producing as many as 500 drafts. She wrote eight hours per day despite a growing list of health problems, including arthritis so severe that she had difficulty even sitting at her typewriter or turning on her desk lamp. Many of these poems never were finalized, and only a few were published in her lifetime.


title: Djuna Barnes

text: During her Patchin Place years, Barnes became a notorious recluse, intensely suspicious of anyone she did not know well. E.E. Cummings, who lived across the street, checked on her periodically by shouting out his window "Are you still alive, Djuna?" Bertha Harris put roses in her mailbox, but never succeeded in meeting her; Carson McCullers camped on her doorstep, but Barnes only called down "Whoever is ringing this bell, please go the hell away." Anaïs Nin was an ardent fan of her work, especially "Nightwood". She wrote to Barnes several times, inviting her to participate in a journal on women's writing, but received no reply. Barnes remained contemptuous of Nin and would cross the street to avoid her. Barnes was angry that Nin had named a character Djuna, and when the feminist bookstore Djuna Books opened in Greenwich Village, Barnes called to demand that the name be changed. Barnes had a lifelong affection for poet Marianne Moore since she and Moore were young in the 1920s.


title: Djuna Barnes

text: Although Barnes had other female lovers, in her later years she was known to claim "I am not a lesbian; I just loved Thelma."


title: Djuna Barnes

text: Barnes was elected to the National Institute of Arts and Letters in 1961 and was awarded a senior fellowship by the National Endowment for the Arts in 1981.


title: Djuna Barnes

text: Barnes was the last surviving member of the first generation of English-language modernists when she died in her home in New York on June 18, 1982, six days after her 90th birthday.


title: Djuna Barnes

text: Barnes's chapbook "The Book of Repulsive Women" (1915) collects eight "rhythms" and five drawings. The poems show the strong influence of late 19th century Decadence, and the style of the illustrations resembles Aubrey Beardsley's. The setting is New York City, and the subjects are all women: a cabaret singer, a woman seen through an open window from the elevated train, and, in the last poem, the corpses of two suicides in the morgue. The book describes women's bodies and sexuality in terms that have indeed struck many readers as repulsive, but, as with much of Barnes's work, the author's stance is ambiguous. Some critics read the poems as exposing and satirizing cultural attitudes toward women.


title: Djuna Barnes

text: Barnes came to regard "The Book of Repulsive Women" as an embarrassment; she called the title "idiotic," left it out of her curriculum vitae, and even burned copies. But because the copyright had never been registered, she was unable to prevent it from being republished, and it became one of her most reprinted works.


title: Djuna Barnes

text: Barnes's novel "Ryder" (1928) draws heavily on her childhood experiences in Cornwall-on-Hudson. It covers 50 years of history of the Ryder family: Sophia Grieve Ryder, like Zadel a former salon hostess fallen into poverty; her idle son Wendell; his wife Amelia; his resident mistress Kate-Careless; and their children. Barnes appears as Wendell and Amelia's daughter Julie. The story has a large cast and is told from a variety of points of view; some characters appear as the protagonist of a single chapter only to disappear from the text entirely. Fragments of the Ryder family chronicle are interspersed with children's stories, songs, letters, poems, parables, and dreams. The book changes style from chapter to chapter, parodying writers from Chaucer to Dante Gabriel Rossetti.


title: Djuna Barnes

text: Both "Ryder" and "Ladies Almanack" abandon the Beardsleyesque style of her drawings for "The Book of Repulsive Women" in favor of a visual vocabulary borrowed from French folk art. Several illustrations are closely based on the engravings and woodcuts collected by Pierre Louis Duchartre and René Saulnier in the 1926 book "L'Imagerie Populaire"—images that had been copied with variations since medieval times. The bawdiness of "Ryder"'s illustrations led the U.S. Postal Service to refuse to ship it, and several had to be left out of the first edition, including an image in which Sophia is seen urinating into a chamberpot and one in which Amelia and Kate-Careless sit by the fire knitting codpieces. Parts of the text were also expurgated. In an acerbic introduction, Barnes explained that the missing words and passages had been replaced with asterisks so that readers could see the "havoc" wreaked by censorship. A 1990 Dalkey Archive edition restored the missing drawings, but the original text was lost with the destruction of the manuscript in World War II.


title: Djuna Barnes

text: "Ladies Almanack" (1928) is a roman à clef about a predominantly lesbian social circle centering on Natalie Clifford Barney's salon in Paris. It is written in an archaic, Rabelaisian style, with Barnes's own illustrations in the style of Elizabethan woodcuts.


title: Djuna Barnes

text: Clifford Barney appears as Dame Evangeline Musset, "who was in her Heart one Grand Red Cross for the Pursuance, the Relief and the Distraction, of such Girls as in their Hinder Parts, and their Fore Parts, and in whatsoever Parts did suffer them most, lament Cruelly." "[A] Pioneer and a Menace" in her youth, Dame Musset has reached "a witty and learned Fifty"; she rescues women in distress, dispenses wisdom, and upon her death is elevated to sainthood. Also appearing pseudonymously are Élisabeth de Gramont, Romaine Brooks, Dolly Wilde, Radclyffe Hall and her partner Una Troubridge, Janet Flanner and Solita Solano, and Mina Loy.


title: Djuna Barnes

text: The obscure language, inside jokes, and ambiguity of "Ladies Almanack" have kept critics arguing about whether it is an affectionate satire or a bitter attack, but Barnes loved the book and reread it throughout her life.


title: Djuna Barnes

text: Barnes's reputation as a writer was made when "Nightwood" was published in England in 1936 in an expensive edition by Faber and Faber, and in America in 1937 by Harcourt, Brace and Company, with an added introduction by T.S. Eliot, Barnes's editor. The avant-garde novel, written under the patronage of Peggy Guggenheim, made Barnes famous in feminist circles.


title: Djuna Barnes

text: The novel, set in Paris in the 1920s, revolves around the lives of five characters, two of whom are based on Barnes and Wood, and it reflects the circumstances surrounding the ending of their relationship. In his introduction, Eliot praises Barnes's style, which, while having "prose rhythm ... and the musical pattern which is not that of verse, is so good a novel that only sensibilities trained on poetry can wholly appreciate it."


title: Djuna Barnes

text: Due to concerns about censorship, Eliot edited "Nightwood" to soften some language relating to sexuality and religion. An edition restoring these changes, edited by Cheryl J. Plumb, was published by Dalkey Archive Press in 1995.


title: Djuna Barnes

text: Dylan Thomas described "Nightwood" as "one of the three great prose books ever written by a woman," and William Burroughs called it "one of the great books of the twentieth century." It was number 12 on a list of the top 100 gay books compiled by The Publishing Triangle in 1999.


title: Djuna Barnes

text: Barnes's verse play "The Antiphon" (1958) is set in England in 1939. Jeremy Hobbs, in disguise as Jack Blow, has brought his family together at their ruined ancestral home, Burley Hall. His motive is never explicitly stated, but he seems to want to provoke a confrontation among the members of his family and force them to confront the truth about their past. His sister Miranda is a stage actress, now "out of patron and of money"; her materialistic brothers, Elisha and Dudley, see her as a threat to their financial well-being. Elisha and Dudley accuse their mother Augusta of complicity with their abusive father Titus Hobbs. They take advantage of Jeremy's absence to don animal masks and assault both women, making cruel and sexually suggestive remarks; Augusta treats this attack as a game. Jeremy returns with a doll house, a miniature version of the house in America where the children grew up. As she examines it, he charges her with making herself "a madam by submission" because she failed to prevent Titus from orchestrating Miranda's rape by "a travelling Cockney thrice [her] age." The last act finds Miranda and Augusta alone together. Augusta, at once disapproving and envious of her daughter's more liberated life, exchanges clothes with her daughter and wants to pretend she is young again, but Miranda refuses to enter into this play. When Augusta hears Elisha and Dudley driving away, she blames Miranda for their abandonment and beats her to death with a curfew bell, falling dead at her side from the exertion.


title: Djuna Barnes

text: The play premiered in 1961 in Stockholm in a Swedish translation by Karl Ragnar Gierow and U.N. Secretary-General Dag Hammarskjöld.


title: Djuna Barnes

text: The play was translated in French and set at the Odeon Theater in Paris by the “Comedie Française” in March 1990.


title: Djuna Barnes

text: Barnes's last book, "Creatures in an Alphabet" (1982), is a collection of short rhyming poems. The format suggests a children's book, but it contains enough allusiveness and advanced vocabulary to make it an unlikely read for a child: the entry for T quotes Blake's "The Tyger," a seal is compared to Jacques-Louis David's portrait of Madame Récamier, and a braying donkey is described as "practicing "solfeggio"." "Creatures" continues the themes of nature and culture found in Barnes's earlier work, and their arrangement as a bestiary reflects her longstanding interest in systems for organizing knowledge, such as encyclopedias and almanacs.


title: Djuna Barnes

text: During the 1920s and 1930s, Barnes made an effort to write a biography of Elsa von Freytag-Loringhoven and edited her poems for publication. Unsuccessful in finding a publisher for the Baroness's poetry, Barnes abandoned the project. After writing a number of chapter drafts of the biography, she abandoned that project, too, after submitting the first chapter to Emily Coleman in 1939, whose response was not encouraging. Barnes's efforts at writing the biography are detailed in Irene Gammel's "Baroness Elsa" biography.


title: Djuna Barnes

text: Barnes has been cited as an influence by writers as diverse as Truman Capote, William Goyen, Karen Blixen, John Hawkes, Bertha Harris, Dylan Thomas, David Foster Wallace, and Anaïs Nin. Writer Bertha Harris described her work as "practically the only available expression of lesbian culture we have in the modern western world" since Sappho.


title: Djuna Barnes

text: Barnes's biographical notes and collection of manuscripts have been a major source for scholars who have brought the Baroness Elsa von Freytag-Loringhoven forth from the margins of Dada history. They were key in producing "" (2011), the first major English collection of the baroness's poems, and a biography titled "Baroness Elsa: Gender, Dada and Everyday Modernity" (2002).


title: Djuna Barnes

text: Cynthia Grant and Svetlana Zylin co-wrote the play "Djuna: What of the Night" based on Barnes's life and works. The play premiered in 1991.


title: Djuna Barnes

text: Emmanuelle Uzan played Barnes in a brief cameo role with no dialogue in Woody Allen's 2011 film "Midnight in Paris".


title: Actin

text: Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over 100 μM; its mass is roughly 42 kDa, with a diameter of 4 to 7 nm.


title: Actin

text: An actin protein is the monomeric subunit of two types of filaments in cells: microfilaments, one of the three major components of the cytoskeleton, and thin filaments, part of the contractile apparatus in muscle cells. It can be present as either a free monomer called G-actin (globular) or as part of a linear polymer microfilament called F-actin (filamentous), both of which are essential for such important cellular functions as the mobility and contraction of cells during cell division.


title: Actin

text: Actin participates in many important cellular processes, including muscle contraction, cell motility, cell division and cytokinesis, vesicle and organelle movement, cell signaling, and the establishment and maintenance of cell junctions and cell shape. Many of these processes are mediated by extensive and intimate interactions of actin with cellular membranes. In vertebrates, three main groups of actin isoforms, alpha, beta, and gamma have been identified. The alpha actins, found in muscle tissues, are a major constituent of the contractile apparatus. The beta and gamma actins coexist in most cell types as components of the cytoskeleton, and as mediators of internal cell motility. It is believed that the diverse range of structures formed by actin enabling it to fulfill such a large range of functions is regulated through the binding of tropomyosin along the filaments.


title: Actin

text: A cell's ability to dynamically form microfilaments provides the scaffolding that allows it to rapidly remodel itself in response to its environment or to the organism's internal signals, for example, to increase cell membrane absorption or increase cell adhesion in order to form cell tissue. Other enzymes or organelles such as cilia can be anchored to this scaffolding in order to control the deformation of the external cell membrane, which allows endocytosis and cytokinesis. It can also produce movement either by itself or with the help of molecular motors. Actin therefore contributes to processes such as the intracellular transport of vesicles and organelles as well as muscular contraction and cellular migration. It therefore plays an important role in embryogenesis, the healing of wounds, and the invasivity of cancer cells. The evolutionary origin of actin can be traced to prokaryotic cells, which have equivalent proteins. Actin homologs from prokaryotes and archaea polymerize into different helical or linear filaments consisting of one or multiple strands. However the in-strand contacts and nucleotide binding sites are preserved in prokaryotes and in archaea. Lastly, actin plays an important role in the control of gene expression.


title: Actin

text: A large number of illnesses and diseases are caused by mutations in alleles of the genes that regulate the production of actin or of its associated proteins. The production of actin is also key to the process of infection by some pathogenic microorganisms. Mutations in the different genes that regulate actin production in humans can cause muscular diseases, variations in the size and function of the heart as well as deafness. The make-up of the cytoskeleton is also related to the pathogenicity of intracellular bacteria and viruses, particularly in the processes related to evading the actions of the immune system.


title: Actin

text: Actin's primary role in the cell is to form linear polymers called microfilaments that serve various functions in the cell's structure, trafficking networks, migration, and replication. The multifaceted role of actin relies on a few of the microfilaments' properties: First, the formation of actin filaments is reversible, and their function often involves undergoing rapid polymerization and depolymerization. Second, microfilaments are polarized – i.e. the two ends of a filament are distinct from one another. Third, actin filaments can bind to many other proteins, which together help modify and organize microfilaments for their diverse functions.


title: Actin

text: Actin is extremely abundant in most cells, comprising 1–5% of the total protein mass of most cells, and 10% of muscle cells.


title: Actin

text: The actin protein is found in both the cytoplasm and the cell nucleus. Its location is regulated by cell membrane signal transduction pathways that integrate the stimuli that a cell receives stimulating the restructuring of the actin networks in response.


title: Actin

text: There are a number of different types of actin with slightly different structures and functions. α-actin is found exclusively in muscle fibres, while β- and γ-actin are found in other cells. As the latter types have a high turnover rate the majority of them are found outside permanent structures. Microfilaments found in cells other than muscle cells are present in three forms:


title: Actin

text: Actin's cytoskeleton is key to the processes of endocytosis, cytokinesis, determination of cell polarity and morphogenesis in yeasts. In addition to relying on actin, these processes involve 20 or 30 associated proteins, which all have a high degree of evolutionary conservation, along with many signalling molecules. Together these elements allow a spatially and temporally modulated assembly that defines a cell's response to both internal and external stimuli.


title: Actin

text: Yeasts contain three main elements that are associated with actin: patches, cables, and rings. Despite not being present for long, these structures are subject to a dynamic equilibrium due to continual polymerization and depolymerization. They possess a number of accessory proteins including ADF/cofilin, which has a molecular weight of 16kDa and is coded for by a single gene, called "COF1"; Aip1, a cofilin cofactor that promotes the disassembly of microfilaments; Srv2/CAP, a process regulator related to adenylate cyclase proteins; a profilin with a molecular weight of approximately 14 kDa that is related/associated with actin monomers; and twinfilin, a 40 kDa protein involved in the organization of patches.


title: Actin

text: Plant genome studies have revealed the existence of protein isovariants within the actin family of genes. Within "Arabidopsis thaliana", a model organism, there are ten types of actin, six profilins, and dozens of myosins. This diversity is explained by the evolutionary necessity of possessing variants that slightly differ in their temporal and spatial expression. The majority of these proteins were jointly expressed in the tissue analysed. Actin networks are distributed throughout the cytoplasm of cells that have been cultivated "in vitro". There is a concentration of the network around the nucleus that is connected via spokes to the cellular cortex, this network is highly dynamic, with a continuous polymerization and depolymerization.


title: Actin

text: Even though the majority of plant cells have a cell wall that defines their morphology, their microfilaments can generate sufficient force to achieve a number of cellular activities, such as the cytoplasmic currents generated by the microfilaments and myosin. Actin is also involved in the movement of organelles and in cellular morphogenesis, which involve cell division as well as the elongation and differentiation of the cell.


title: Actin

text: The most notable proteins associated with the actin cytoskeleton in plants include: villin, which belongs to the same family as gelsolin/severin and is able to cut microfilaments and bind actin monomers in the presence of calcium cations; fimbrin, which is able to recognize and unite actin monomers and which is involved in the formation of networks (by a different regulation process from that of animals and yeasts); formins, which are able to act as an F-actin polymerization nucleating agent; myosin, a typical molecular motor that is specific to eukaryotes and which in "Arabidopsis thaliana" is coded for by 17 genes in two distinct classes; CHUP1, which can bind actin and is implicated in the spatial distribution of chloroplasts in the cell; KAM1/MUR3 that define the morphology of the Golgi apparatus as well as the composition of xyloglucans in the cell wall; NtWLIM1, which facilitates the emergence of actin cell structures; and ERD10, which is involved in the association of organelles within membranes and microfilaments and which seems to play a role that is involved in an organism's reaction to stress.


title: Actin

text: Nuclear actin was first noticed and described in 1977 by Clark and Merriam. Authors describe a protein present in the nuclear fraction, obtained from "Xenopus laevis" oocytes, which shows the same features as skeletal muscle actin. Since that time there have been many scientific reports about the structure and functions of actin in the nucleus (for review see: Hofmann 2009.) The controlled level of actin in the nucleus, its interaction with actin-binding proteins (ABP) and the presence of different isoforms allows actin to play an important role in many important nuclear processes.


title: Actin

text: The actin sequence does not contain a nuclear localization signal. The small size of actin (about 43 kDa) allows it to enter the nucleus by passive diffusion. The import of actin into the nucleus (probably in a complex with cofilin) is facilitated by the import protein importin 9.


title: Actin

text: Low levels of actin in the nucleus seems to be important, because actin has two nuclear export signals (NES) in its sequence. Microinjected actin is quickly removed from the nucleus to the cytoplasm. Actin is exported at least in two ways, through exportin 1 and exportin 6. Specific modifications, such as SUMOylation, allows for nuclear actin retention. A mutation preventing SUMOylation causes rapid export of beta actin from the nucleus.


title: Actin

text: Nuclear actin exists mainly as a monomer, but can also form dynamic oligomers and short polymers. Nuclear actin organization varies in different cell types. For example, in "Xenopus" oocytes (with higher nuclear actin level in comparison to somatic cells) actin forms filaments, which stabilize nucleus architecture. These filaments can be observed under the microscope thanks to fluorophore-conjugated phalloidin staining.


title: Actin

text: In somatic cell nuclei, however, actin filaments cannot be observed using this technique. The DNase I inhibition assay, the only test which allows the quantification of the polymerized actin directly in biological samples, has revealed that endogenous nuclear actin indeed occurs mainly in a monomeric form.


title: Actin

text: Precisely controlled level of actin in the cell nucleus, lower than in the cytoplasm, prevents the formation of filaments. The polymerization is also reduced by the limited access to actin monomers, which are bound in complexes with ABPs, mainly cofilin.


title: Actin

text: Different isoforms of actin are present in the cell nucleus. The level of actin isoforms may change in response to stimulation of cell growth or arrest of proliferation and transcriptional activity. Research on nuclear actin is focused on isoform beta. However the use of antibodies directed against different actin isoforms allows identifying not only the cytoplasmic beta in the cell nucleus, but also alpha- and gamma-actin in certain cell types. The presence of different isoforms of actin may have a significant effect on its function in nuclear processes, as the level of individual isoforms can be controlled independently.


title: Actin

text: Functions of actin in the nucleus are associated with its ability to polymerize and interact with various ABPs and with structural elements of the nucleus. Nuclear actin is involved in:


title: Actin

text: Due to its ability to undergo conformational changes and interaction with many proteins, actin acts as a regulator of formation and activity of protein complexes such as transcriptional complex.


title: Actin

text: Actin is also involved in cell movement. A meshwork of actin filaments marks the forward edge of a moving cell, and the polymerization of new actin filaments pushes the cell membrane forward in protrusions called lamellipodia. These membrane protrusions then attach to the substrate, forming structures known as focal adhesions that connect to the actin network. Once attached, the rear of the cell body contracts squeezing its contents forward past the adhesion point. Once the adhesion point has moved to the rear of the cell, the cell disassembles it, allowing the rear of the cell to move forward.


title: Actin

text: In addition to the physical force generated by actin polymerization, microfilaments facilitate the movement of various intracellular components by serving as the roadway along which a family of motor proteins called myosins travel.


title: Actin

text: Actin plays a particularly prominent role in muscle cells, which consist largely of repeated bundles of actin and myosin II. Each repeated unit – called a sarcomere – consists of two sets of oppositely oriented F-actin strands ("thin filaments"), interlaced with bundles of myosin ("thick filaments"). The two sets of actin strands are oriented with their (+) ends embedded in either end of the sarcomere in delimiting structures called Z-disks. The myosin fibrils are in the middle between the sets of actin filaments, with strands facing in both directions. When the muscle contracts, the myosin threads move along the actin filaments towards the (+) end, pulling the ends of the sarcomere together and shortening it by around 70% of its length. In order to move along the actin thread, myosin must hydrolyze ATP; thus ATP serves as the energy source for muscle contraction.


title: Actin

text: At times of rest, the proteins tropomyosin and troponin bind to the actin filaments, preventing the attachment of myosin. When an activation signal (i.e. an action potential) arrives at the muscle fiber, it triggers the release of Ca from the sarcoplasmic reticulum into the cytosol. The resulting spike in cytosolic calcium rapidly releases tropomyosin and troponin from the actin thread, allowing myosin to bind, and muscle contracation to begin.


title: Actin

text: In the final stages of cell division, many cells form a ring of actin at the cell's midpoint. This ring, aptly called the "contractile ring", uses a similar mechanism as muscle fibers where myosin II pulls along the actin ring, causing it to contract. This contraction cleaves the parent cell into two, completing cytokinesis. The contractile ring is composed of actin, myosin, anillin, and α-actinin. In the fission yeast "Schizosaccharomyces pombe", actin is actively formed in the constricting ring with the participation of Arp3, the formin Cdc12, profilin, and WASp, along with preformed microfilaments. Once the ring has been constructed the structure is maintained by a continual assembly and disassembly that, aided by the Arp2/3 complex and formins, is key to one of the central processes of cytokinesis.


title: Actin

text: Actin-myosin pairs can also participate in the trafficking of various membrane vesicles and organelles within the cell. Myosin V is activated by binding to various cargo receptors on organelles, and then moves along an actin filament towards the (+) end, pulling its cargo along with it.


title: Actin

text: These nonconventional myosins use ATP hydrolysis to transport cargo, such as vesicles and organelles, in a directed fashion much faster than diffusion. Myosin V walks towards the barbed end of actin filaments, while myosin VI walks toward the pointed end. Most actin filaments are arranged with the barbed end toward the cellular membrane and the pointed end toward the cellular interior. This arrangement allows myosin V to be an effective motor for the export of cargos, and myosin VI to be an effective motor for import.


title: Actin

text: The traditional image of actin's function relates it to the maintenance of the cytoskeleton and, therefore, the organization and movement of organelles, as well as the determination of a cell's shape. However, actin has a wider role in eukaryotic cell physiology, in addition to similar functions in prokaryotes.


title: Actin

text: Monomeric actin, or G-actin, has a globular structure consisting of two lobes separated by a deep cleft. The bottom of the cleft represents the “ATPase fold”, a structure conserved among ATP and GTP-binding proteins that binds to a magnesium ion and a molecule of ATP. Binding of ATP or ADP is required to stabilize each actin monomer; without one of these molecules bound, actin quickly becomes denatured.


title: Actin

text: The X-ray crystallography model of actin that was produced by Kabsch from the striated muscle tissue of rabbits is the most commonly used in structural studies as it was the first to be purified. The G-actin crystallized by Kabsch is approximately 67 x 40 x 37 Å in size, has a molecular mass of 41,785 Da and an estimated isoelectric point of 4.8. Its net charge at pH = 7 is -7.


title: Actin

text: Elzinga and co-workers first determined the complete peptide sequence for this type of actin in 1973, with later work by the same author adding further detail to the model. It contains 374 amino acid residues. Its N-terminus is highly acidic and starts with an acetyled aspartate in its amino group. While its C-terminus is alkaline and is formed by a phenylalanine preceded by a cysteine, which has a degree of functional importance. Both extremes are in close proximity within the I-subdomain. An anomalous "N"-methylhistidine is located at position 73.


title: Actin

text: The tertiary structure is formed by two domains known as the large and the small, which are separated by a cleft centred around the location of the bond with ATP-ADP+P. Below this there is a deeper notch called a “groove”. In the native state, despite their names, both have a comparable depth.


title: Actin

text: The normal convention in topological studies means that a protein is shown with the biggest domain on the left-hand side and the smallest domain on the right-hand side. In this position the smaller domain is in turn divided into two: subdomain I (lower position, residues 1–32, 70–144, and 338–374) and subdomain II (upper position, residues 33–69). The larger domain is also divided in two: subdomain III (lower, residues 145–180 and 270–337) and subdomain IV (higher, residues 181–269). The exposed areas of subdomains I and III are referred to as the “barbed” ends, while the exposed areas of domains II and IV are termed the “pointed" ends. This nomenclature refers to the fact that, due to the small mass of subdomain II actin is polar; the importance of this will be discussed below in the discussion on assembly dynamics. Some authors call the subdomains Ia, Ib, IIa, and IIb, respectively.


title: Actin

text: The most notable supersecondary structure is a five chain beta sheet that is composed of a β-meander and a β-α-β clockwise unit. It is present in both domains suggesting that the protein arose from gene duplication.


title: Actin

text: Under various conditions, G-actin molecules polymerize into longer threads called "filamentous-" or "F-actin". These F-actin threads are typically composed of two helical strands of actin wound around each other, forming a 7 to 9 nanometer wide helix that repeats every 72 nanometers (or every 14 G-actin subunits). In F-actin threads, G-actin molecules are all oriented in the same direction. The two ends of the F-actin thread are distinct from one another. At one end – designated the (−) end – the ATP-binding cleft of the terminal actin molecule is facing outward. At the opposite end – designated (+) – the ATP-binding cleft is buried in the filament, contacting the neighboring actin molecule. As F-actin threads grow, new molecules tend to join at the (+) end of an existing F-actin strand. Conversely, threads tend to shrink by shedding actin monomers from the strand's (-) end.


title: Actin

text: Some proteins, such as cofilin appear to increase the angle of turn, but again this could be interpreted as the establishment of different structural states. These could be important in the polymerization process.


title: Actin

text: There is less agreement regarding measurements of the turn radius and filament thickness: while the first models assigned a length of 25 Å, current X-ray diffraction data, backed up by cryo-electron microscopy suggests a length of 23.7 Å. These studies have shown the precise contact points between monomers. Some are formed with units of the same chain, between the "barbed" end on one monomer and the "pointed" end of the next one. While the monomers in adjacent chains make lateral contact through projections from subdomain IV, with the most important projections being those formed by the C-terminus and the hydrophobic link formed by three bodies involving residues 39–42, 201–203, and 286. This model suggests that a filament is formed by monomers in a "sheet" formation, in which the subdomains turn about themselves, this form is also found in the bacterial actin homologue MreB.


title: Actin

text: The terms "pointed" and "barbed" referring to the two ends of the microfilaments derive from their appearance under transmission electron microscopy when samples are examined following a preparation technique called "decoration". This method consists of the addition of myosin S1 fragments to tissue that has been fixed with tannic acid. This myosin forms polar bonds with actin monomers, giving rise to a configuration that looks like arrows with feather fletchings along its shaft, where the shaft is the actin and the fletchings are the myosin. Following this logic, the end of the microfilament that does not have any protruding myosin is called the point of the arrow (- end) and the other end is called the barbed end (+ end).


title: Actin

text: A S1 fragment is composed of the head and neck domains of myosin II. Under physiological conditions, G-actin (the monomer form) is transformed to F-actin (the polymer form) by ATP, where the role of ATP is essential.


title: Actin

text: The helical F-actin filament found in muscles also contains a tropomyosin molecule, which is a 40 nanometre long protein that is wrapped around the F-actin helix. During the resting phase the tropomyosin covers the actin's active sites so that the actin-myosin interaction cannot take place and produce muscular contraction. There are other protein molecules bound to the tropomyosin thread, these are the troponins that have three polymers: troponin I, troponin T, and troponin C.


title: Actin

text: F-actin is both strong and dynamic. Unlike other polymers, such as DNA, whose constituent elements are bound together with covalent bonds, the monomers of actin filaments are assembled by weaker bonds. The lateral bonds with neighbouring monomers resolve this anomaly, which in theory should weaken the structure as they can be broken by thermal agitation. In addition, the weak bonds give the advantage that the filament ends can easily release or incorporate monomers. This means that the filaments can be rapidly remodelled and can change cellular structure in response to an environmental stimulus. Which, along with the biochemical mechanism by which it is brought about is known as the "assembly dynamic".


title: Actin

text: Actin can spontaneously acquire a large part of its tertiary structure. However, the way it acquires its fully functional form from its newly synthesized native form is special and almost unique in protein chemistry. The reason for this special route could be the need to avoid the presence of incorrectly folded actin monomers, which could be toxic as they can act as inefficient polymerization terminators. Nevertheless, it is key to establishing the stability of the cytoskeleton, and additionally, it is an essential process for coordinating the cell cycle.


title: Actin

text: CCT is required in order to ensure that folding takes place correctly. CCT is a group II chaperonin, a large protein complex that assists in the folding of other proteins. CCT is formed of a double ring of eight different subunits (hetero-octameric) and it differs from group I chaperonins like GroEL, which is found in Eubacteria and in eukaryotic organelles, as it does not require a co-chaperone to act as a lid over the central catalytic cavity. Substrates bind to CCT through specific domains. It was initially thought that it only bound with actin and tubulin, although recent immunoprecipitation studies have shown that it interacts with a large number of polypeptides, which possibly function as substrates. It acts through ATP-dependent conformational changes that on occasion require several rounds of liberation and catalysis in order to complete a reaction.


title: Actin

text: In order to successfully complete their folding, both actin and tubulin need to interact with another protein called prefoldin, which is a heterohexameric complex (formed by six distinct subunits), in an interaction that is so specific that the molecules have coevolved. Actin complexes with prefoldin while it is still being formed, when it is approximately 145 amino acids long, specifically those at the N-terminal.


title: Actin

text: Different recognition sub-units are used for actin or tubulin although there is some overlap. In actin the subunits that bind with prefoldin are probably PFD3 and PFD4, which bind in two places one between residues 60–79 and the other between residues 170–198. The actin is recognized, loaded, and delivered to the cytosolic chaperonin (CCT) in an open conformation by the inner end of prefoldin's "tentacles” (see the image and note). The contact when actin is delivered is so brief that a tertiary complex is not formed, immediately freeing the prefoldin.


title: Actin

text: The CCT then causes actin's sequential folding by forming bonds with its subunits rather than simply enclosing it in its cavity. This is why it possesses specific recognition areas in its apical β-domain. The first stage in the folding consists of the recognition of residues 245–249. Next, other determinants establish contact. Both actin and tubulin bind to CCT in open conformations in the absence of ATP. In actin's case, two subunits are bound during each conformational change, whereas for tubulin binding takes place with four subunits. Actin has specific binding sequences, which interact with the δ and β-CCT subunits or with δ-CCT and ε-CCT. After AMP-PNP is bound to CCT the substrates move within the chaperonin's cavity. It also seems that in the case of actin, the CAP protein is required as a possible cofactor in actin's final folding states.


title: Actin

text: The exact manner by which this process is regulated is still not fully understood, but it is known that the protein PhLP3 (a protein similar to phosducin) inhibits its activity through the formation of a tertiary complex.


title: Actin

text: Actin is an ATPase, which means that it is an enzyme that hydrolyzes ATP. This group of enzymes is characterised by their slow reaction rates. It is known that this ATPase is “active”, that is, its speed increases by some 40,000 times when the actin forms part of a filament. A reference value for this rate of hydrolysis under ideal conditions is around 0.3 s. Then, the P remains bound to the actin next to the ADP for a long time, until it is cooperatively liberated from the interior of the filament.


title: Actin

text: The exact molecular details of the catalytic mechanism are still not fully understood. Although there is much debate on this issue, it seems certain that a "closed" conformation is required for the hydrolysis of ATP, and it is thought that the residues that are involved in the process move to the appropriate distance. The glutamic acid Glu137 is one of the key residues, which is located in subdomain 1. Its function is to bind the water molecule that produces a nucleophilic attack on the ATP's γ-phosphate bond, while the nucleotide is strongly bound to subdomains 3 and 4. The slowness of the catalytic process is due to the large distance and skewed position of the water molecule in relation to the reactant. It is highly likely that the conformational change produced by the rotation of the domains between actin's G and F forms moves the Glu137 closer allowing its hydrolysis. This model suggests that the polymerization and ATPase's function would be decoupled straight away. The "open" to "closed" transformation between G and F forms and its implications on the relative motion of several key residues and the formation of water wires have been characterized in molecular dynamics and QM/MM simulations.


title: Actin

text: Actin filaments are often rapidly assembled and disassembled, allowing them to generate force and support cell movement. Assembly classically occurs in three steps. First, the "nucleation phase", in which two to three G-actin molecules slowly join to form a small oligomer that will nucleate further growth. Second, the "elongation phase", when the actin filament rapidly grows by the addition of many actin molecules to both ends. As the filament grows, actin molecules are added to the (+) end of the filament around 10 times faster than to the (−) end, and so filaments tend to primarily grow at the (+) end. Third, the "steady-state phase", where an equillibrium is reached as actin molecules join and leave the filament at the same rate, maintaining the filament's length. While the filament's length remains constant in the steady-state phase, new molecules are constantly being added to the (+) end and falling off the (−) end, a phenomenon called "treadmilling" as a given actin molecule would appear to move along the strand. In isolation, whether a filament will grow or shrink, and how quickly, are determined by the concentration of G-actin around the filament; however, in cells, the dynamics of actin filaments are heavily influenced by various actin-binding proteins.


title: Actin

text: The actin cytoskeleton "in vivo" is not exclusively composed of actin, other proteins are required for its formation, continuance, and function. These proteins are called actin-binding proteins and they are involved in actin's polymerization, depolymerization, stability, and organisation. The diversity of these proteins is such that actin is thought to be the protein that takes part in the greatest number of protein-protein interactions.


title: Actin

text: The nucleation of new actin filaments – the rate-limiting step in actin polymerization – is aided by actin-nucleating proteins such as formins (like formin-2) and the Arp2/3 complex. Formins help to nucleate long actin filaments. They bind two free actin-ATP molecules, bringing them together. Then as the filament begins to grow, formin moves along the (+) end of the growing filament, all the while recruiting actin-binding proteins that promote filament growth, and excluding capping proteins that would block filament extension. Branches in actin filaments are typically nucleated by the Arp2/3 complex in concert with nucleation promoting factors. Nucleation promoting factors bind two free G-actin molecules, then recruit and activate the Arp2/3 complex. The activated Arp2/3 complex attaches to an existing actin filament, and uses the two bound G-actin molecules to nucleate a new actin filament branching off of the old one at a 70° angle.


title: Actin

text: As filaments grow, the pool of available G-actin molecules is managed by G-actin-binding proteins such as profilin and thymosin β-4. Profilin ensures a supply of available actin-ATP by binding to ADP-bound G-actin and promoting the exchange of ADP for ATP. Profilin's binding to the actin molecule physically blocks its addition to a filament's (−) end, but permits it to join the (+) end. Once the actin-ATP has joined the filament, profilin releases it. As formins promote the nucleation and extension of new actin filaments, they recruit profilin to the area, increasing the local concentration of actin-ATP to boost filament growth. In contrast, thymosin β-4 binds and sequesters actin-ATP, preventing it from joining a microfilament.


title: Actin

text: Once an actin fiber is established, the dynamics of its growth or collapse are influenced by numerous proteins. Existing strands can be interrupted by filament cleaving proteins, such as cofilin and gelsolin. Cofilin binds along two actin-ADP molecules in a filament, forcing a movement that destabilizes the filament and causes it to break. Gelsolin inserts itself between actin molecules in a filament, disrupting the filament. After the filament breaks, gelsolin remains attached to the new (+) end, preventing it from growing, thus forcing its disassembly.


title: Actin

text: Other proteins bind to the ends of actin filaments, stabilizing them. These are called "capping proteins" and include CapZ and tropomodulin. CapZ binds the (+) end of a filament, preventing further addition or loss of actin from that end. Tropomodulin binds to a filament's (−) end, again preventing addition or loss of molecule's at that end. Tropomodulin is typically found in cells that require extremely stable actin filaments, such as those in muscle and red blood cells.


title: Actin

text: These actin binding proteins are typically regulated by various cellular signals to control actin assembly dynamics in different cellular locations. Formins, for example, are typically folded in an inactive conformation until they're activated by the binding of the small GTPase Rho. Actin branching at the cell membrane is important for cell movement, and so the plasma membrane lipid PIP activates the nucleation promoting factor WASp and inhibits CapZ. WASp is also activated by the small GTPase Cdc42, while another nucleation promoting factor WAVE is activated by the GTPase Rac1.


title: Actin

text: Although most yeasts have only a single actin gene, higher eukaryotes, in general, express several isoforms of actin encoded by a family of related genes. Mammals have at least six actin isoforms coded by separate genes, which are divided into three classes – alpha, beta, and gamma – according to their isoelectric points. In general, alpha actins are found in muscle (α-skeletal, α-aortic smooth, α-cardiac), whereas beta and gamma isoforms are prominent in non-muscle cells (β-cytoplasmic, γ1-cytoplasmic, γ2-enteric smooth). Although the amino acid sequences and "in vitro" properties of the isoforms are highly similar, these isoforms cannot completely substitute for one another "in vivo". Plants contains more than 60 actin genes and pseudogenes.


title: Actin

text: The typical actin gene has an approximately 100-nucleotide 5' UTR, a 1200-nucleotide translated region, and a 200-nucleotide 3' UTR. The majority of actin genes are interrupted by introns, with up to six introns in any of 19 well-characterised locations. The high conservation of the family makes actin the favoured model for studies comparing the introns-early and introns-late models of intron evolution.


title: Actin

text: Actin and closely related proteins are present in all organisms, suggesting the common ancestor of all life on Earth had actin. Actin is one of the most conserved proteins throughout the evolution of eukaryotes. The sequences of actin proteins from animals and amoebae are 80% identical despite being separated by approximately one billion years of evolution. Many unicellular eukaryotes have a single actin gene, while multicellular eukaryotes often have several closely related genes that serve specialized functions. Humans have six; plants have 10 or more. In addition to actin, eukaryotes have a large family of actin-related proteins, or "Arps", that share a common ancestor with actin and are called Arp1–Arp11, with Arp1 the most closely related to actin, and Arp11 the least.


title: Actin

text: Bacteria encode three types of actin: MreB influences cell shape, FtsA cell division, and ParM separation of large plasmids. Some archaea have a bacteria-like MreB gene, while others have an actin gene that more closely resembles eukaryote actin.


title: Actin

text: The eukaryotic cytoskeleton of organisms among all taxonomic groups have similar components to actin and tubulin. For example, the protein that is coded by the "ACTG2" gene in humans is completely equivalent to the homologues present in rats and mice, even though at a nucleotide level the similarity decreases to 92%. However, there are major differences with the equivalents in prokaryotes (FtsZ and MreB), where the similarity between nucleotide sequences is between 40 and 50 % among different bacteria and archaea species. Some authors suggest that the ancestral protein that gave rise to the model eukaryotic actin resembles the proteins present in modern bacterial cytoskeletons.


title: Actin

text: Some authors point out that the behaviour of actin, tubulin, and histone, a protein involved in the stabilization and regulation of DNA, are similar in their ability to bind nucleotides and in their ability of take advantage of Brownian motion. It has also been suggested that they all have a common ancestor. Therefore, evolutionary processes resulted in the diversification of ancestral proteins into the varieties present today, conserving, among others, actins as efficient molecules that were able to tackle essential ancestral biological processes, such as endocytosis.


title: Actin

text: The bacterial cytoskeleton contains proteins that are highly similar to actin monomers and polymers. The bacterial protein MreB polymerizes into thin non-helical filaments and occasionally into helical structures similar to F-actin. Furthermore, its crystalline structure is very similar to that of G-actin (in terms of its three-dimensional conformation), there are even similarities between the MreB protofilaments and F-actin. The bacterial cytoskeleton also contains the FtsZ proteins, which are similar to tubulin.


title: Actin

text: Bacteria therefore possess a cytoskeleton with homologous elements to actin (for example, MreB, AlfA, ParM, FtsA, and MamK), even though the amino acid sequence of these proteins diverges from that present in animal cells. However, such proteins have a high degree of structural similarity to eukaryotic actin. The highly dynamic microfilaments formed by the aggregation of MreB and ParM are essential to cell viability and they are involved in cell morphogenesis, chromosome segregation, and cell polarity. ParM is an actin homologue that is coded in a plasmid and it is involved in the regulation of plasmid DNA. ParMs from different bacterial plasmids can form astonishingly diverse helical structures comprising two or four strands to maintain faithful plasmid inheritance.


title: Actin

text: The majority of mammals possess six different actin genes. Of these, two code for the cytoskeleton ("ACTB" and "ACTG1") while the other four are involved in skeletal striated muscle ("ACTA1"), smooth muscle tissue ("ACTA2"), intestinal muscles ("ACTG2") and cardiac muscle ("ACTC1"). The actin in the cytoskeleton is involved in the pathogenic mechanisms of many infectious agents, including HIV. The vast majority of the mutations that affect actin are point mutations that have a dominant effect, with the exception of six mutations involved in nemaline myopathy. This is because in many cases the mutant of the actin monomer acts as a “cap” by preventing the elongation of F-actin.


title: Actin

text: "ACTA1" is the gene that codes for the α-isoform of actin that is predominant in human skeletal striated muscles, although it is also expressed in heart muscle and in the thyroid gland. Its DNA sequence consists of seven exons that produce five known transcripts. The majority of these consist of point mutations causing substitution of amino acids. The mutations are in many cases associated with a phenotype that determines the severity and the course of the affliction.


title: Actin

text: The mutation alters the structure and function of skeletal muscles producing one of three forms of myopathy: type 3 nemaline myopathy, congenital myopathy with an excess of thin myofilaments (CM) and congenital myopathy with fibre type disproportion (CMFTD). Mutations have also been found that produce core myopathies. Although their phenotypes are similar, in addition to typical nemaline myopathy some specialists distinguish another type of myopathy called actinic nemaline myopathy. In the former, clumps of actin form instead of the typical rods. It is important to state that a patient can show more than one of these phenotypes in a biopsy. The most common symptoms consist of a typical facial morphology (myopathic facies), muscular weakness, a delay in motor development and respiratory difficulties. The course of the illness, its gravity, and the age at which it appears are all variable and overlapping forms of myopathy are also found. A symptom of nemaline myopathy is that "nemaline rods" appear in differing places in type 1 muscle fibres. These rods are non-pathognomonic structures that have a similar composition to the Z disks found in the sarcomere.


title: Actin

text: The pathogenesis of this myopathy is very varied. Many mutations occur in the region of actin's indentation near to its nucleotide binding sites, while others occur in Domain 2, or in the areas where interaction occurs with associated proteins. This goes some way to explain the great variety of clumps that form in these cases, such as Nemaline or Intranuclear Bodies or Zebra Bodies. Changes in actin's folding occur in nemaline myopathy as well as changes in its aggregation and there are also changes in the expression of other associated proteins. In some variants where intranuclear bodies are found the changes in the folding masks the nucleus's protein exportation signal so that the accumulation of actin's mutated form occurs in the cell nucleus. On the other hand, it appears that mutations to "ACTA1" that give rise to a CFTDM have a greater effect on sarcomeric function than on its structure. Recent investigations have tried to understand this apparent paradox, which suggests there is no clear correlation between the number of rods and muscular weakness. It appears that some mutations are able to induce a greater apoptosis rate in type II muscular fibres.


title: Actin

text: "ACTG2" codes for the largest actin isoform, which has nine exons, one of which, the one located at the 5' end, is not translated. It is a γ-actin that is expressed in the enteric smooth muscle. No mutations to this gene have been found that correspond to pathologies, although microarrays have shown that this protein is more often expressed in cases that are resistant to chemotherapy using cisplatin.


title: Actin

text: "ACTA2" codes for an α-actin located in the smooth muscle, and also in vascular smooth muscle. It has been noted that the MYH11 mutation could be responsible for at least 14% of hereditary thoracic aortic aneurisms particularly Type 6. This is because the mutated variant produces an incorrect filamentary assembly and a reduced capacity for vascular smooth muscle contraction. Degradation of the aortic media has been recorded in these individuals, with areas of disorganization and hyperplasia as well as stenosis of the aorta's vasa vasorum. The number of afflictions that the gene is implicated in is increasing. It has been related to Moyamoya disease and it seems likely that certain mutations in heterozygosis could confer a predisposition to many vascular pathologies, such as thoracic aortic aneurysm and ischaemic heart disease. The α-actin found in smooth muscles is also an interesting marker for evaluating the progress of liver cirrhosis.


title: Actin

text: The "ACTC1" gene codes for the α-actin isoform present in heart muscle. It was first sequenced by Hamada and co-workers in 1982, when it was found that it is interrupted by five introns. It was the first of the six genes where alleles were found that were implicated in pathological processes.


title: Actin

text: A number of structural disorders associated with point mutations of this gene have been described that cause malfunctioning of the heart, such as Type 1R dilated cardiomyopathy and Type 11 hypertrophic cardiomyopathy. Certain defects of the atrial septum have been described recently that could also be related to these mutations.


title: Actin

text: Two cases of dilated cardiomyopathy have been studied involving a substitution of highly conserved amino acids belonging to the protein domains that bind and intersperse with the Z discs. This has led to the theory that the dilation is produced by a defect in the transmission of contractile force in the myocytes.


title: Actin

text: The mutations in ACTC1 are responsible for at least 5% of hypertrophic cardiomyopathies. The existence of a number of point mutations have also been found:


title: Actin

text: Pathogenesis appears to involve a compensatory mechanism: the mutated proteins act like toxins with a dominant effect, decreasing the heart's ability to contract causing abnormal mechanical behaviour such that the hypertrophy, that is usually delayed, is a consequence of the cardiac muscle's normal response to stress.


title: Actin

text: Recent studies have discovered ACTC1 mutations that are implicated in two other pathological processes: Infantile idiopathic restrictive cardiomyopathy, and noncompaction of the left ventricular myocardium.


title: Actin

text: "ACTB" is a highly complex locus. A number of pseudogenes exist that are distributed throughout the genome, and its sequence contains six exons that can give rise to up to 21 different transcriptions by alternative splicing, which are known as the β-actins. Consistent with this complexity, its products are also found in a number of locations and they form part of a wide variety of processes (cytoskeleton, NuA4 histone-acyltransferase complex, cell nucleus) and in addition they are associated with the mechanisms of a great number of pathological processes (carcinomas, juvenile dystonia, infection mechanisms, nervous system malformations and tumour invasion, among others). A new form of actin has been discovered, kappa actin, which appears to substitute for β-actin in processes relating to tumours.


title: Actin

text: Three pathological processes have so far been discovered that are caused by a direct alteration in gene sequence:


title: Actin

text: The "ACTG1" locus codes for the cytosolic γ-actin protein that is responsible for the formation of cytoskeletal microfilaments. It contains six exons, giving rise to 22 different mRNAs, which produce four complete isoforms whose form of expression is probably dependent on the type of tissue they are found in. It also has two different DNA promoters. It has been noted that the sequences translated from this locus and from that of β-actin are very similar to the predicted ones, suggesting a common ancestral sequence that suffered duplication and genetic conversion.


title: Actin

text: In terms of pathology, it has been associated with processes such as amyloidosis, retinitis pigmentosa, infection mechanisms, kidney diseases, and various types of congenital hearing loss.


title: Actin

text: Six autosomal-dominant point mutations in the sequence have been found to cause various types of hearing loss, particularly sensorineural hearing loss linked to the DFNA 20/26 locus. It seems that they affect the stereocilia of the ciliated cells present in the inner ear's Organ of Corti. β-actin is the most abundant protein found in human tissue, but it is not very abundant in ciliated cells, which explains the location of the pathology. On the other hand, it appears that the majority of these mutations affect the areas involved in linking with other proteins, particularly actomyosin. Some experiments have suggested that the pathological mechanism for this type of hearing loss relates to the F-actin in the mutations being more sensitive to cofilin than normal.


title: Actin

text: However, although there is no record of any case, it is known that γ-actin is also expressed in skeletal muscles, and although it is present in small quantities, model organisms have shown that its absence can give rise to myopathies.


title: Actin

text: Some infectious agents use actin, especially cytoplasmic actin, in their life cycle. Two basic forms are present in bacteria:


title: Actin

text: In addition to the previously cited example, actin polymerization is stimulated in the initial steps of the internalization of some viruses, notably HIV, by, for example, inactivating the cofilin complex.


title: Actin

text: Actin is used in scientific and technological laboratories as a track for molecular motors such as myosin (either in muscle tissue or outside it) and as a necessary component for cellular functioning. It can also be used as a diagnostic tool, as several of its anomalous variants are related to the appearance of specific pathologies.


title: Actin

text: The use of actin as an internal control is based on the assumption that its expression is practically constant and independent of experimental conditions. By comparing the expression of the gene of interest to that of the actin, it is possible to obtain a relative quantity that can be compared between different experiments, whenever the expression of the latter is constant. It is worth pointing out that actin does not always have the desired stability in its gene expression.


title: Actin

text: Actin was first observed experimentally in 1887 by W.D. Halliburton, who extracted a protein from muscle that 'coagulated' preparations of myosin that he called "myosin-ferment". However, Halliburton was unable to further refine his findings, and the discovery of actin is credited instead to Brunó Ferenc Straub, a young biochemist working in Albert Szent-Györgyi's laboratory at the Institute of Medical Chemistry at the University of Szeged, Hungary.


title: Actin

text: Following up on the discovery of Ilona Banga & Szent-Györgyi in 1941 that the coagulation only occurs in some myosin extractions and was reversed upon the addition of ATP, Straub identified and purified actin from those myosin preparations that did coagulate. Building on Banga's original extraction method, he developed a novel technique for extracting muscle protein that allowed him to isolate substantial amounts of relatively pure actin, published in 1942. Straub's method is essentially the same as that used in laboratories today. Since Straub's protein was necessary to activate the coagulation of myosin, it was dubbed "actin". Realizing that Banga's coagulating myosin preparations contained actin as well, Szent-Györgyi called the mixture of both proteins actomyosin.


title: Actin

text: The hostilities of World War II meant Szent-Gyorgyi was unable to publish his lab's work in Western scientific journals. Actin therefore only became well known in the West in 1945, when their paper was published as a supplement to the "Acta Physiologica Scandinavica". Straub continued to work on actin, and in 1950 reported that actin contains bound ATP and that, during polymerization of the protein into microfilaments, the nucleotide is hydrolyzed to ADP and inorganic phosphate (which remain bound to the microfilament). Straub suggested that the transformation of ATP-bound actin to ADP-bound actin played a role in muscular contraction. In fact, this is true only in smooth muscle, and was not supported through experimentation until 2001.


title: Actin

text: The amino acid sequencing of actin was completed by M. Elzinga and co-workers in 1973. The crystal structure of G-actin was solved in 1990 by Kabsch and colleagues. In the same year, a model for F-actin was proposed by Holmes and colleagues following experiments using co-crystallization with different proteins. The procedure of co-crystallization with different proteins was used repeatedly during the following years, until in 2001 the isolated protein was crystallized along with ADP. However, there is still no high-resolution X-ray structure of F-actin. The crystallization of G-actin was possible due to the use of a rhodamine conjugate that impedes polymerization by blocking the amino acid cys-374. Christine Oriol-Audit died in the same year that actin was first crystallized but she was the researcher that in 1977 first crystallized actin in the absence of Actin Binding Proteins (ABPs). However, the resulting crystals were too small for the available technology of the time.


title: Actin

text: Although no high-resolution model of actin's filamentous form currently exists, in 2008 Sawaya's team were able to produce a more exact model of its structure based on multiple crystals of actin dimers that bind in different places. This model has subsequently been further refined by Sawaya and Lorenz. Other approaches such as the use of cryo-electron microscopy and synchrotron radiation have recently allowed increasing resolution and better understanding of the nature of the interactions and conformational changes implicated in the formation of actin filaments.


title: Actin

text: A number of natural toxins that interfere with actin's dynamics are widely used in research to study actin's role in biology. Latrunculin – a toxin produced by sponges – binds to G-actin preventing it from joining microfilaments. Cytochalasin D – produced by certain fungi – serves as a capping factor, binding to the (+) end of a filament and preventing further addition of actin molecules. In contrast, the sponge toxin jasplakinolide promotes the nucleation of new actin filaments by binding and stabilzing pairs of actin molecules. Phalloidin – from the "death cap" mushroom "Amanita phalloides" – binds to adjacent actin molecules within the F-actin filament, stabilizing the filament and preventing its depolymerization.


title: Actin

text: Phalloidin is often labelled with fluorescent dyes to visualize actin filaments by fluorescence microscopy.


title: The Murder of Nicole Brown Simpson

text: The Murder of Nicole Brown Simpson is a 2019 American crime horror film directed by Daniel Farrands. The film is loosely based on the murder of Nicole Brown Simpson, presenting a version of events in which Brown Simpson is murdered by serial killer Glen Edward Rogers, and not by O. J. Simpson, her ex-husband and the primary suspect in the case. Though Mena Suvari's performance as Nicole Brown was praised, the film was universally panned by critics.


title: The Murder of Nicole Brown Simpson

text: Review aggregator Rotten Tomatoes gives the film a approval rating, based on reviews, with an average rating of . Frank Scheck of "The Hollywood Reporter" gave the film a negative review, stating that it (along with several of the director's other films) was the equivalent of "cinematic graverobbing" and that one of the film's only fans will likely be O. J. Simpson. Guy Lodge of "Variety" also gave it a negative review, calling it an "unabashedly tacky true-crime thriller" and writing that "it's a cheap, unloving death march of a movie", though Lodge offered some praise to Mena Suvari in the title role.


title: Sun Dance

text: The Sun Dance is a ceremony practiced by some Native Americans in the United States and Indigenous peoples in Canada, primarily those of the Plains cultures. It usually involves the community gathering together to pray for healing. Individuals make personal sacrifices on behalf of the community.


title: Sun Dance

text: After European colonization of the Americas, and with the formation of the Canadian and United States governments, both countries passed laws intended to suppress Indigenous cultures and force assimilation to majority-European culture.


title: Sun Dance

text: The Sun Dance was one of the prohibited ceremonies, as was the potlatch of the Pacific Northwest peoples. Canada lifted its prohibition against the practice of the full ceremony in 1951. In the United States, Congress passed the American Indian Religious Freedom Act (AIRFA) in 1978, which was enacted to protect basic civil liberties, and to protect and preserve the traditional religious rights and cultural practices of Native Americans, Eskimos, Aleuts, and Native Hawaiians.


title: Sun Dance

text: Several features are common to the ceremonies held by Sun Dance cultures. These include dances and songs passed down through many generations, the use of a traditional drum, a sacred fire, praying with a ceremonial pipe, fasting from food and water before participating in the dance, and, in some cases, the ceremonial piercing of skin and a trial of physical endurance. Certain plants are picked and prepared for use during the ceremony.


title: Sun Dance

text: Typically, the Sun Dance is a grueling ordeal for the dancers, a physical and spiritual test that they offer in sacrifice for their people. According to the Oklahoma Historical Society, young men dance around a pole to which they are fastened by "rawhide thongs pegged through the skin of their chests."


title: Sun Dance

text: While not all Sun Dance ceremonies include piercing, the object of the Sun Dance is to offer personal sacrifice for the benefit of one's family and community. The dancers fast for many days, in the open air and whatever weather occurs.


title: Sun Dance

text: At most ceremonies, family members and friends stay in the surrounding camp and pray in support of the dancers. Much time and energy by the entire community are needed to conduct the sun dance gatherings and ceremonies. Communities plan and organize for at least a year to prepare for the ceremony. Usually, one leader or a small group of leaders are in charge of the ceremony, but many elders help out and advise. A group of helpers do many of the tasks required to prepare for the ceremony.


title: Sun Dance

text: The Government of Canada, through the Department of Indian Affairs (now Crown-Indigenous Relations and Northern Development Canada), persecuted Sun Dance practitioners and attempted to suppress the dance. Indian agents, based on directives from their superiors, did routinely interfere with, discouraged, and disallowed sun dances on many Canadian plains communities from 1882 until the 1940s.


title: Sun Dance

text: The Canadian government outlawed "any celebration or dance of which the wounding or mutilation of the dead or living body of any human being or animal forms a part or is a feature" in an 1895 amendment to the Indian Act. Anyone who engaged, assisted or encouraged (either directly or indirectly) was liable to imprisonment. Though not all nations' Sun Dances include the body piercings, the amendment legally prohibited its performance for those communities that did.


title: Sun Dance

text: It is unclear how often this law was actually enforced; in at least one instance, police are known to have given their permission for the ceremony to be conducted. The First Nations people simply conducted many ceremonies quietly and in secret. Sun dance practitioners, such as the Plains Cree, Saulteaux, and Blackfoot, continued to hold sun dances throughout the persecution period. Some practiced the dance in secret, others with permissions from their agents, and others without the body piercing.


title: Sun Dance

text: In 1951, government officials amended the "Indian Act", dropping the prohibition against practices of flesh-wounding.


title: Sun Dance

text: The Sun Dance is practiced annually in many First Nations communities in Canada. The Cree and Saulteaux have conducted at least one Rain Dance (with similar elements) each year since 1880 somewhere on the Canadian Plains.


title: Sun Dance

text: In 1993, responding to what they believed was a frequent desecration of the Sun Dance and other Lakota sacred ceremonies, US and Canadian Lakota, Dakota and Nakota nations held "the Lakota Summit V". It was an international gathering of about 500 representatives from 40 different peoples and bands of the Lakota. They unanimously passed the following 'Declaration of War Against Exploiters of Lakota Spirituality':


title: Sun Dance

text: "Whereas sacrilegious "sundances" for non-Indians are being conducted by charlatans and cult leaders who promote abominable and obscene imitations of our sacred Lakota sundance rites; ... We hereby and henceforth declare war against all persons who persist in exploiting, abusing, and misrepresenting the sacred traditions and spiritual practices of the Lakota, Dakota and Nakota people." - Mesteth, Wilmer, et al (1993)


title: Sun Dance

text: In 1995, efforts to continue practice the ceremony on an tract of unceded Secwepemc land led to an armed confrontation known as the Gustafsen Lake standoff.


title: Sun Dance

text: In 2003, the 19th-Generation Keeper of the Sacred White Buffalo Calf Pipe of the Lakota asked non-Indigenous people to stop attending the Sun Dance ("Wi-wayang-wa-c'i-pi" in Lakota); he stated that all can pray in support, but that only Indigenous people should approach the altars. This statement was supported by keepers of sacred bundles and traditional spiritual leaders from the Cheyenne, Dakota, Lakota, and Nakota nations, who issued a proclamation that non-Indigenous people would be banned from sacred altars and the Seven Sacred Rites, including and especially the sun dance, effective March 9, 2003 onward:


title: Sun Dance

text: In most Sun Dance cultures, it is forbidden to film ceremony or prayer. Few images exist of authentic ceremonies. Many First Nations people believe that when money or cameras enter, the spirits leave, so no photo conveys an authentic ceremony. The Kainai Nation in Alberta permitted filming of their Sun Dance in the late 1950s. This was released as the documentary "Circle of the Sun" (1960), produced by the National Film Board of Canada. Manitoba archival photos show that the ceremonies have been consistently practiced since at least the early 1900s.


title: Studebaker Avanti

text: The Studebaker Avanti is a personal luxury coupe manufactured and marketed by Studebaker Corporation between June 1962 and December 1963. A halo car for the maker, it was marketed as "America's only four-passenger high-performance personal car."


title: Studebaker Avanti

text: Described as "one of the more significant milestones of the postwar industry", the Raymond Loewy-designed car offered safety features and high-speed performance. Called “the fastest production car in the world” upon its introduction, a modified Avanti reached over with its supercharged R3 engine at the Bonneville Salt Flats. In all, it broke 29 world speed records at the Bonneville Salt Flats.


title: Studebaker Avanti

text: Following Studebaker's discontinuation of the model, a series of five owner arrangements continued to manufacture and market derivatives of the Avanti model through 2006.


title: Studebaker Avanti

text: The Avanti was developed at the direction of Studebaker president, Sherwood Egbert, who took over in February 1961. The car's design theme was "allegedly doodled by Egbert on the proverbial back of an envelope during an airplane flight." Egbert's 'doodle' was to answer Ford's Thunderbird and an attempt to improve the automaker's sagging performance. Designed by Raymond Loewy's team of Tom Kellogg, Bob Andrews, and John Ebstein on a 40-day crash program, the Avanti featured a radical fiberglass body mounted on a modified Studebaker Lark 109-inch convertible chassis and powered by a modified 289 Hawk engine. A Paxton supercharger was offered as an option.


title: Studebaker Avanti

text: In eight days the stylists finished a "clay scale model with two different sides: one a two-place sports car, the other a four-seat GT coupe." Tom Kellogg, a young California stylist hired for this project by Loewy, "felt it should be a four-seat coupe." "Loewy envisioned a low-slung, long-hood-short-deck semi-fastback coupe with a grilleless nose and a wasp-waisted curvature to the rear fenders, suggesting a supersonic aircraft."


title: Studebaker Avanti

text: The Avanti's complex body shape "would have been both challenging and prohibitively expensive to build in steel" with Studebaker electing to mold the exterior panels in glass-reinforced plastic (fiberglass), outsourcing the work to Molded Fiberglass Body in Ashtabula, Ohio — the same company that built the fiberglass panels for the Chevrolet Corvette in 1953.


title: Studebaker Avanti

text: The Avanti featured front disc-brakes that were British Dunlop designed units, made under license by Bendix, "the first American production model to offer them." It was one of the first bottom breather designs where air enters from under the front of the vehicle rather than via a conventional grille, a design feature much more common after the 1980s.


title: Studebaker Avanti

text: The Avanti was publicly introduced on April 26, 1962, "simultaneously at the New York International Automobile Show and at the Annual Shareholders' Meeting." Rodger Ward, winner of the 1962 Indianapolis 500, received a Studebaker Avanti as part of his prize package, "thus becoming the first private owner of an Avanti." A Studebaker Lark convertible was the Indianapolis pace car that year and the Avanti was named the honorary pace car.


title: Studebaker Avanti

text: In December 1962 the "Los Angeles Times" reported: "Launching of operations at Studebaker's own fiber-glass body works to increase the production of Avantis." Many production problems concerning the supplier, fit, and finish resulted in delays and cancelled orders.


title: Studebaker Avanti

text: After the closure of Studebaker's factory on December 20, 1963, "Competition Press" reported: "Avantis will no longer be manufactured and contrary to the report that there are thousands gathering dust in South Bend warehouses, Studebaker has only five Avantis left. Dealers have about 2,500, and 1,600 have been sold since its introduction." This contrasted with Chevrolet which produced 23,631 Corvette sports cars in 1963. According to the book "My Father The Car" written about Stu Chapman, Studebaker Corporation's Advertising & Public Relations Department head in Canada, Studebaker seriously considered re-introducing the Avanti into Studebaker showrooms in 1965/66 after production resumed in 1965 via Studebaker-Packard dealership owners Newman & Altman.


title: Studebaker Avanti

text: The Avanti name, tooling, and plant space were sold to two South Bend, Indiana, Studebaker dealers, Nate Altman and Leo Newman. They re-introduced a slightly modified hand-built version of the original Avanti using leftover Studebaker chassis and engines from General Motors. There was no connection with the Studebaker brand name.


title: Studebaker Avanti

text: Following Altman and Newman's effort, a succession of additional entrepreneurs purchased the tooling and name to manufacture small numbers of increasingly modified variants of the car, including the Avanti II, through 2006.


title: Studebaker Avanti

text: The Avanti Owners Association International is an active association with nearly 2,000 members worldwide and meeting yearly in various cities in the United States and in Switzerland. Members of the not-for-profit organization receive the full-color quarterly "Avanti Magazine" publication, published since the organization's founding in 1965.


title: Electorate of Bavaria

text: The Electorate of Bavaria () was an independent hereditary electorate of the Holy Roman Empire from 1623 to 1806, when it was succeeded by the Kingdom of Bavaria.


title: Electorate of Bavaria

text: The Wittelsbach dynasty which ruled the Duchy of Bavaria was the younger branch of the family which also ruled the Electorate of the Palatinate. The head of the elder branch was one of the seven prince-electors of the Holy Roman Empire according to the Golden Bull of 1356, but Bavaria was excluded from the electoral dignity. In 1621, the Elector Palatine Frederick V was put under the imperial ban for his role in the Bohemian Revolt against Emperor Ferdinand II, and the electoral dignity and territory of the Upper Palatinate was conferred upon his loyal cousin, Duke Maximilian I of Bavaria. Although the Peace of Westphalia would create a new electoral title for Frederick V's son, with the exception of a brief period during the War of the Spanish Succession, Maximilian's descendants would continue to hold the original electoral dignity until the extinction of his line in 1777. At that point the two lines were joined in personal union until the end of the Holy Roman Empire. In 1805, after the Peace of Pressburg, the then-elector, Maximilian Joseph, raised himself to the dignity of King of Bavaria, and the Holy Roman Empire was abolished the year after.