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COPI
COPI
from Wikipedia
Electron micrograph of in vitro–formed COPI-coated vesicles. Average vesicle diameter at the membrane level is 60 nm.

COPI is a coatomer, a protein complex[1] that coats vesicles transporting proteins from the cis end of the Golgi complex back to the rough endoplasmic reticulum (ER), where they were originally synthesized, and between Golgi compartments. This type of transport[clarification needed] is retrograde transport, in contrast to the anterograde transport associated with the COPII protein. The name "COPI" refers to the specific coat protein complex that initiates the budding process on the cis-Golgi membrane. The coat consists of large protein subcomplexes that are made of seven different protein subunits, namely α, β, β', γ, δ, ε and ζ.

COPI
Identifiers
SymbolCOPI_C
PfamPF06957
InterProIPR010714
Available protein structures:
PDB  IPR010714 PF06957 (ECOD; PDBsum)  
AlphaFold

Coat proteins

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Coat protein, or COPI, is an ADP ribosylation factor (ARF)-dependent protein involved in membrane traffic.[2] COPI was first identified in retrograde traffic from the cis-Golgi to the rough endoplasmic reticulum (ER)[3][4] and is the most extensively studied of ARF-dependent adaptors. COPI consists of seven subunits which compose the heteroheptameric protein complex.

The primary function of adaptors is the selection of cargo proteins for their incorporation into nascent carriers. Cargo containing the sorting motifs KKXX and KXKXX interact with COPI to form carriers which are transported from the cis-Golgi to the ER.[5][6][7][8][9] Current views suggest that ARFs are also involved in the selection of cargo for incorporation into carriers.

Budding process

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ADP ribosylation factor (ARF) is a GTPase involved in membrane traffic. There are 6 mammalian ARFs which are regulated by over 30 guanine nucleotide exchange factors (GEFs) and GTPase activating proteins (GAPs). ARF is post-translationally modified at the N-terminus by the addition of the fatty acid myristate.

ARF cycles between GTP and GDP-bound conformations. In the GTP-bound form, ARF conformation changes such that the myristate and hydrophobic N-terminal become more exposed and associate with the membrane. The interconversion between GTP and GDP bound states is mediated by ARF GEFs and ARF GAPs. At the membrane, ARF-GTP is hydrolyzed to ARF-GDP by ARF GAPs. Once in the GDP-bound conformation, ARF converts to a less hydrophobic conformation and dissociates from the membrane. Soluble ARF-GDP is converted back to ARF-GTP by GEFs.

  1. Luminal proteins: Proteins found in the lumen of the Golgi complex that need to be transported to the lumen of the ER contain the signal peptide KDEL.[10] This sequence is recognized by a membrane-bound KDEL receptor. In yeast, this is ERD2P and in mammals it is KDELR. This receptor then binds to an ARF-GEF, a class of guanine nucleotide exchange factors. This protein in turn binds to the ARF. This interaction causes ARF to exchange its bound GDP for GTP. Once this exchange is made ARF binds to the cytosolic side of the cis-Golgi membrane and inserts the myristoylated N-terminal amphipathic alpha-helix into the membrane.[11]
  2. Membrane proteins: Transmembrane proteins which reside in the ER contain sorting signals in their cytosolic tails which direct the protein to exit the Golgi and return to the ER. These sorting signals, or motifs, typically contain the amino acid sequence KKXX or KXKXX, which interact with COPI subunits α-COP and β'-COP.[10][9] The order in which adaptor proteins associate with cargo, or adaptor proteins associate with ARFs is unclear, however, in order to form a mature transport carrier coat protein, adaptor, cargo, and ARF must all associate.

Membrane deformation and carrier budding occurs following the collection of interactions described above. The carrier then buds off of the donor membrane, in the case of COPI this membrane is the cis-Golgi, and the carrier moves to the ER where it fuses with the acceptor membrane and its content is expelled.

Structure

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The COPI triad. Color scheme: membrane - gray; Arf1 - pink; gamma-COP - light green; beta-COP, dark green; zeta-COP - yellow; delta-COP - orange; betaprime-COP - light blue; alpha-COP - dark blue

On the surface of a vesicle COPI molecules form symmetric trimers ("triads"). The curved triad structure positions the Arf1 molecules and cargo binding sites proximal to the membrane. The β′- and α-COP subunits form an arch over the γζβδ-COP subcomplex, orienting their N-terminal domains such that the K(X)KXX cargo-motif binding sites are optimally positioned against the membrane. Thus β′- and α-COP do not form a cage or lattice as in COPII and clathrin coats as previously suggested;[12] instead, they are linked to one another via the γζβδ-COP subcomplexes, forming an interconnected assembly.[13] The triads are linked together with contacts of variable valence making up four different types of contacts.[14]

See also

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References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Copi is an Argentine cartoonist, playwright, and novelist known for his minimalist drawing style, sharp satirical humor, and provocative explorations of sexuality, gender, and power in both comics and theater. Born Raúl Damonte Botana in Buenos Aires on November 20, 1939, he adopted the pseudonym Copi (derived from "copito de nieve," the Spanish diminutive for "little snowflake") and relocated to Paris in 1962, where he spent most of his career. His cartoons, characterized by simple lines and absurd, often transgressive scenarios, appeared in prominent French publications such as Le Nouvel Observateur and Libération, establishing him as a key figure in underground and alternative press. As a playwright, Copi authored experimental works that blended camp aesthetics, black humor, and queer themes, including pieces like Eva Perón, The Homosexual, or the Difficulty of Expressing Oneself, and Loretta Strong, many of which he also performed in, frequently in drag. Copi's literary output extended to novels such as The Uruguayan and The Queens' Ball, which further showcased his irreverent style and critique of bourgeois society. An openly gay artist during a time of limited visibility, his work significantly influenced queer literature and performance art in France and Latin America. He died in Paris on December 14, 1987, from AIDS-related complications, leaving a legacy as one of the most original and boundary-pushing voices in 20th-century humor and theater.

Early Life

Family Background and Childhood

Copi was born Raúl Damonte Botana on November 20, 1939, in Buenos Aires, Argentina, into a family prominent in journalism and politics. His pseudonym "Copi" originated from his grandmother's nickname for him, "copita de nieve" (little snowflake). His maternal grandfather was Natalio Félix Botana, a major figure in Argentine journalism who founded the influential newspaper Crítica and supported numerous intellectuals and artists. His father, Raúl Damonte Taborda, was a journalist, politician affiliated with the Radical party, and director of the newspaper Tribuna Popular, known for his anti-Peronist stance after initially supporting Perón before breaking with him and conducting campaigns against Peronism from exile. Due to his father's political activities and opposition to Juan Perón, the family endured multiple periods of exile, including stays in Uruguay, Haiti, and New York City. Copi's youth was primarily spent in Montevideo, Uruguay, which he later described as a kind of lost paradise.

Early Artistic Beginnings in Argentina

Copi demonstrated a precocious talent for drawing from an early age. During his adolescence, he began contributing caricatures to Tribuna Popular, the newspaper directed by his father, as well as to the satirical magazine Tía Vicenta. These early works marked the start of his engagement with graphic humor and satire in the Latin American context, though they remained occasional rather than full-time professional endeavors before his emigration in 1962. In the 1950s, Copi produced his first humorous cartoons for Tía Vicenta, where he created the notable character La mujer sentada. His developing style drew significant influence from the French cartoonist Jean Bosc, whose graphic approach shaped Copi's emerging talent for witty and satirical illustration. These contributions reflected the vibrant satirical press scene in Argentina at the time and laid the foundation for his later distinctive minimalist humor.

Emigration to France

Relocation to Paris in 1962

In 1962, Copi permanently relocated to Paris at the age of 23, marking his final emigration from Argentina to France, where he would live until his death in 1987. This move shifted his artistic career from its early development in the Latin American context to the dynamic French and European avant-garde environment. Upon arriving in Paris, Copi began contributing to French magazines such as Tweety and Bizarre, establishing an initial foothold in the local publishing scene. He also engaged in early collaborations with the Franco-Argentine artists' association Tse, connecting with expatriate and local creative networks. During the 1960s, Copi participated in the avant-garde Panic movement, working alongside Alejandro Jodorowsky, Fernando Arrabal, and Roland Topor in experimental and provocative artistic endeavors. This involvement reflected his rapid adaptation to the radical and interdisciplinary spirit of the Parisian intellectual and artistic circles.

Integration into the French Avant-Garde Scene

Copi quickly became involved with the Panic Movement in the 1960s, a transgressive artistic collective founded by Fernando Arrabal, Alejandro Jodorowsky, and Roland Topor, though some sources describe him as joining or associating closely with its founders rather than co-founding. He collaborated with these figures on experimental performances and happenings that emphasized shock, absurdity, and anti-establishment provocation, aligning with the group's ethos of disrupting conventional artistic norms. His contributions to the satirical and underground press, particularly through cartoons published in Hara-Kiri, further cemented his reputation within Paris's countercultural and avant-garde circles during this period. In 1969, as a member of the Tse association of Franco-Argentine artists, Copi staged his biographical play Eva Perón at the Théâtre de l'Épée de Bois, an event that provoked international shock and controversy for its irreverent and hallucinatory portrayal of the Argentine figure. This production highlighted his growing prominence as an Argentine émigré artist fully embedded in the French avant-garde scene. Through these networks and activities, Copi established himself as a distinctive voice in Paris's experimental artistic landscape of the 1960s, bridging Argentine roots with French underground innovation.

Cartooning Career

Minimalist Style and Influences

Copi's cartooning is renowned for its minimalist style, characterized by extreme simplicity in linework, highly reduced figures that verge on abstraction, and an economical use of visual elements that leaves much to the viewer's imagination. This stripped-down approach amplifies the absurdity in his gags, often juxtaposing mundane or polite scenarios with sudden bursts of grotesque, violent, or morbid content delivered in a deadpan manner. One of his primary graphic influences was the French cartoonist Jean Bosc, whose own extreme minimalism—featuring sparse lines, empty backgrounds, and naïvely rendered characters—served as a model for contrasting innocent appearances with pitch-black satire and sharp social commentary. Copi drew from this French satirical tradition, adapting its irreverent spirit to create provocative, personal narratives that frequently probed taboos and societal hypocrisies through absurd logic and unflinching directness. After relocating to France in 1962, Copi's work shifted toward more explicit and transgressive themes, intensifying the erotic, cruel, and apocalyptic elements that had been present but less pronounced in his earlier Argentine cartoons. This evolution allowed him to push the boundaries of graphic humor, blending graphic restraint with bold content to critique conformity, authority, and conventional morality. Copi's distinctive combination of minimalist aesthetics and provocative absurdity exerted influence on later cartoonists, including Georges Wolinski, who adopted similar techniques for incisive and unconventional satire.

Major Publications and Iconic Series

Copi's cartooning career in France was marked by regular contributions to leading satirical and cultural magazines, where he developed his distinctive voice in humor and social commentary. He published extensively in Hara-Kiri, Charlie Mensuel, Le Nouvel Observateur, and Libération, establishing a presence in the French press from the mid-1960s onward. His drawings also reached international audiences through outlets such as Evergreen in the United States, Linus in Italy, and Triunfo in Spain. Among his most recognized series, Copi launched La Femme assise in 1965 for Le Nouvel Observateur, a recurring feature that became emblematic of his work. The following year, he introduced Les Poulets n'ont pas de chaise in 1966. He contributed to Gai Pied from its founding, serving as part of the magazine's original militant team. Several of Copi's cartoons were compiled into notable collections, including Le dernier salon où l’on cause (1973), Et moi, pourquoi j’ai pas de banane? (1975), Les vieilles putes (1977), and Le monde fantastique des gays (1986). These albums preserved his prolific output from various periodicals and solidified his legacy in French cartooning.

Theatrical Career

Playwriting and Productions

Copi's theatrical career flourished after his arrival in Paris in 1962, where he contributed to the avant-garde scene through his writing, directing, and occasional acting in stage productions. He is credited as the author of works featured in 134 recorded spectacles, directed three of his own plays, and appeared as an actor in seven productions, including some of his own creations. His plays were staged by prominent directors such as Jorge Lavelli, Alfredo Arias, Jérôme Savary, Claude Confortès, Marcial Di Fonzo Bo, and Laurent Pelly, among others who helped disseminate his distinctive voice in French and international theater. He began his playwriting with Un ángel para la señora Lisca, which he directed himself in Buenos Aires in 1962, marking his entry into dramatic writing before his full integration into the French scene. In France, he participated in experimental theater events associated with the Panic Movement, including happenings directed by Jorge Lavelli at Le Bilboquet in 1965 alongside Fernando Arrabal. Many of his works explored themes of failed communication, grotesque exaggeration, and disrupted human interactions, often in a rapid, violent farce style that blended cruelty with dark humor. One of his most notable and controversial plays, Eva Perón, written in 1969, premiered in 1970 at the Théâtre de l'Épée de Bois in Paris under the direction of Alfredo Arias with his TSE troupe. The production featured a male actor in the title role and provoked disruption by extreme right-wing groups offended by its irreverent portrayal of the iconic Argentine figure during her final days battling cancer. The text presents Eva as refusing death, insulting those around her, and indulging in delusions amid absent communication and grotesque power dynamics. Subsequent productions included Laurent Pelly's 1993 staging at Théâtre National de Chaillot and Marcial Di Fonzo Bo's 2001 version in Santiago de Chile, which won the Prix du Syndicat de la critique for best spectacle. Copi directed several of his own works, including La Pyramide in 1975, La Coupe du monde Franzozos in 1977, and Le Frigo in 1984, in which he also performed. Other key plays such as Les Quatre Jumelles premiered in 1973 directed by Jorge Lavelli and have seen numerous revivals over decades. His oeuvre has enjoyed sustained posthumous interest since his death in 1987, with regular stagings and adaptations, including operatic treatments such as L'Ombre de Venceslao in collaboration with Jorge Lavelli and composer Martin Matalon. Copi also took acting roles in films such as Race d'Ep (1979) and The Book of Mary (1985).

Key Plays and Themes

Copi's theatrical works are renowned for their provocative exploration of homosexuality, gender ambiguity, marginality, and absurdity, often through extravagant, grotesque characters who embody sexual uncertainty and bodily decay. His plays feature frequent travestism and delirious, surreal humor, portraying outsiders adrift in existential displacement and cosmic fantasies. These recurring motifs reflect Copi's position as a perpetual marginal figure in both life and art, imposing a form of radical pudeur amid avant-garde excess. His major plays include Sainte Geneviève dans sa baignoire, premiered in 1966, an early experimental piece. La journée d'une rêveuse followed in 1968, depicting a mother constructing her son in a dreamlike narrative. L'homosexuel ou la difficulté de s'exprimer, staged in 1971, centers on a delirious transvestite character obsessed with travel, physical decrepitude, and sexual incertitude, a role Copi performed himself. Les quatre jumelles appeared in 1973, succeeded by Loretta Strong in 1974 and La Pyramide in 1975, each extending his grotesque and satirical vision. Later works sustained these themes: La Tour de la Défense in 1981, a vitriolic comedy, Le Frigo in 1983, La nuit de Madame Lucienne in 1985, and the posthumously premiered Une visite inopportune in 1988, which portrays the final days of a faded actor dying of AIDS in a hospital. These pieces collectively highlight Copi's preoccupation with marginal identities and the absurd limits of human expression.

Literary Career

Novels and Prose Works

Copi's novels and prose works, composed primarily in French with occasional ventures into Spanish, are marked by their provocative, satirical, and transgressive character, often probing themes of identity, sexuality, and societal norms through absurd and grotesque lenses. His narratives feature unstable gender pronouns, queer marginality, and black humor that positions homosexuality as an existential relation to the world rather than a mere identity category. He began his prose career in the 1970s with L'Uruguayen (1973), a luminous and humorous travel narrative drawing on his family's exile from Argentina to Uruguay amid Perón's rise. This was followed by Le bal des folles (1977), a novel recounting a chaotic, decades-spanning love affair across Rome, Paris, New York, and Ibiza amid eccentric figures and the liberated excesses of the era. Une langouste pour deux (1978) collected short stories that subvert banal domestic scenes by unleashing characters' concealed deformities, impulses, and absurdities. His later prose maintained this intensity with La cité des rats (1979), La vida es un tango (1981, his only completed novel written in Spanish), La guerre des pédés (1982), Virginia Woolf a encore frappé (1983), and the posthumous L'Internationale argentine (1988). These works continued to deploy scandalous satire and marginal perspectives to interrogate social conventions and personal identity. His prose shares thematic overlaps with his theatrical output, particularly in its irreverent treatment of queer experience and societal rupture.

Personal Life

Identity, Sexuality, and Recurring Themes

Copi was openly homosexual and immersed himself in Paris's queer artistic and theatrical milieu during the 1970s and 1980s. He was close to the Homosexual Front for Revolutionary Action (FHAR), a key group in the French gay liberation movement, particularly through his relationship as companion of FHAR founder Guy Hocquenghem, and was affiliated with the prominent French gay magazine Gai Pied, reflecting his active engagement with the community and its cultural outlets. His sexual identity profoundly informed recurring motifs across his cartoons, plays, and novels, where themes of homosexuality, gender fluidity, transvestism, and social marginality appeared consistently. Copi employed highly transgressive and surreal approaches to explore the nature of femininity, masculinity, and homosexuality, often presenting shattered identities and kaleidoscopic trans-identities that challenged conventional norms. Through this provocative art, he confronted social taboos surrounding sexuality and gender, using subversion and humor to express queer perspectives.

Death

In his final years, Copi battled an AIDS-related illness that progressively worsened, reflecting the devastating impact of the epidemic during its early years in France. He remained active creatively until shortly before the end, completing works amid declining health. Copi died on December 14, 1987, in Paris at the age of 48 due to complications from AIDS, with contemporary reports citing skin cancer as a direct cause. As one of the prominent cultural figures in France to succumb to the disease in the 1980s, his death highlighted the growing toll of AIDS on the artistic community. Posthumously, his final play Une visite inopportune, a semi-autobiographical comedy depicting a hospital patient dying of AIDS, premiered in February 1988 at the Théâtre de la Colline under Jorge Lavelli's direction and was published that same year. His novel L'Internationale argentine also appeared in 1988. These releases underscored the continuation of his legacy immediately after his passing.

References

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