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Background: The growing importance of the ubiquitous fungal genus Trichoderma (Hypocreales, Ascomycota) requires understanding of its biology and evolution. Many Trichoderma species are used as biofertilizers and biofungicides and T. reesei is the model organism for industrial production of cellulolytic enzymes. In addition, some highly opportunistic species devastate mushroom farms and can become pathogens of humans. A comparative analysis of the first three whole genomes revealed mycoparasitism as the innate feature of Trichoderma. However, the evolution of these traits is not yet understood.
Results: We selected 12 most commonly occurring Trichoderma species and studied the evolution of their genome sequences. Trichoderma evolved in the time of the Cretaceous-Palaeogene extinction event 66 (15) mya, but the formation of extant sections (Longibrachiatum, Trichoderma) or clades (Harzianum/Virens) happened in Oligocene. The evolution of the Harzianum clade and section Trichoderma was accompanied by significant gene gain, but the ancestor of section Longibrachiatum experienced rapid gene loss. The highest number of genes gained encoded ankyrins, HET domain proteins and transcription factors. We also identified the Trichoderma core genome, completely curated its annotation, investigated several gene families in detail and compared the results to those of other fungi. Eighty percent of those genes for which a function could be predicted were also found in other fungi, but only 67% of those without a predictable function.
Conclusions: Our study presents a time scaled pattern of genome evolution in 12 Trichoderma species from three phylogenetically distant clades/sections and a comprehensive analysis of their genes. The data offer insights in the evolution of a mycoparasite towards a generalist.
Manganiello was born in Pittsburgh, Pennsylvania, to Susan and Charles Manganiello.[2] His mother is of German and Armenian descent.[3][4] Manganiello's father was born in Massachusetts outside of Boston.[5] Joe Manganiello was raised in Mt. Lebanon, Pennsylvania.[6]
In the February 7, 2023, episode of PBS series Finding Your Roots, the researchers uncovered that Manganiello's legal paternal grandfather, Emilio Manganiello,[7] was not his biological grandfather, and that his biological great-grandparents were an African-American man named William Henry Cutler and a white woman named Nellie Alton. His biological paternal grandfather was one of Cutler and Alton's three mixed-race African-American sons.[8][5] Using this information, researchers traced Manganiello's paternal lineage back to his great-great-great-great-great-grandfather, Plato Turner, an African slave who was freed before slavery was abolished in Massachusetts and who went on to fight for the Continental Army during the American Revolution.[5][7] Manganiello's maternal great-grandmother, Terviz "Rose" Darakjian,[8] was a survivor of the Armenian genocide, where her husband and seven of her children were killed. Darakjian's eighth child, an infant, drowned during Darakjian's escape across the Euphrates River.[5] Darakjian later encountered Karl Wilhelm Beutinger, a German soldier, in an internment camp for survivors. Rose became pregnant by him. Beutinger soon left to return to Germany. He resumed his life there with his German wife and children, and never saw Rose Darakjian again. Manganiello's grandmother is the child of Beutinger and Rose Darakjian.[8][5][9]
He has a younger brother, Nicholas. Raised Catholic, was a student at St. Bernard School, a Catholic elementary school in Mt. Lebanon, Pennsylvania and then attended Mt. Lebanon High School, where he graduated with honors in 1995 and won the school's Great Alumni Award in 2011.[10][11][12][13][14] Growing up, he was the captain of his football, basketball, and volleyball teams and went on to play at the varsity level in all three sports.[10] He won the role of Jud Fry in his school's senior year production of Oklahoma! and was involved with the school's TV studio. He would borrow equipment in order to write and direct films with his friends, which eventually inspired him to begin studying acting.[13]
Manganiello suffered a series of sports injuries through High School, including a torn medial collateral ligament while returning a kickoff in a varsity football game against Ringgold High School. The time off allowed Manganiello to reevaluate his future and so he decided to audition for the Carnegie Mellon School of Drama during his senior year. He was not accepted, so he enrolled at the University of Pittsburgh and worked over the next year in the theater. He then reapplied to Carnegie Mellon a year later, and was awarded a scholarship, accepted into the acting program that year.[10] There he performed in theater productions and wrote, produced, and acted in a student film entitled Out of Courage 2: Out for Vengeance.[15] He graduated in 2000 with a Bachelor of Fine Arts degree in acting.[13] He then traveled to New York City and Los Angeles through his university to participate in group auditions, which provided him contacts in the entertainment business including an agent, a manager, and a screen test for Sam Raimi's Spider-Man.[16]
While a student at Carnegie Mellon University, Manganiello appeared in numerous productions in Pittsburgh's theatre scene, including Ulfheim in Henrik Ibsen's When We Dead Awaken, Lorenzo in Shakespeare's The Merchant of Venice for Quantum Theatre, and Joe in the Pittsburgh premiere of The Last Night of Ballyhoo.[15] He moved to Los Angeles, California, after graduating from Carnegie Mellon.[16] He quickly signed with a talent agent, and three days later, he auditioned for the role of Peter Parker in the Sam Raimi-directed film Spider-Man (2002). He landed the role of Eugene "Flash" Thompson, Peter Parker's nemesis, as his first acting job out of college.[15] He reprised the role several years later, making a brief cameo in Spider-Man 3 (2007).[17]
Manganiello began finding work in television, playing Tori Spelling's boyfriend on VH1's So Notorious in 2006, and guest starred on Las Vegas, Jake in Progress, and Close to Home. That year, he also played John Leguizamo's Alcoholics Anonymous sponsor in the CBS television pilot Edison. In 2007, he appeared in the Scrubs episode "My No Good Reason" and in the MyNetwork TV nighttime soap opera American Heiress. He also played Officer Litchman, the love interest to Linda Cardellini's character, for a four-episode arc on NBC's ER. He returned to the stage playing The Chick Magnet in May 2007 for the New York City premiere of Skirts & Flirts, a monologue show by Gloria Calderon Kellett, for which he was a finalist for HBO's Aspen Comedy Festival.[18] He then played Stanley Kowalski in A Streetcar Named Desire for the West Virginia Public Theatre in 2008,[19] directed by his former Carnegie Mellon professor Geoffrey Hitch.[18]
He starred as Leo Belraggio, a New York jazz musician, in the West Coast premiere of Terrence McNally's Unusual Acts of Devotion in June 2009. The play was staged at the La Jolla Playhouse at the University of California in San Diego. The summer prior, Manganiello worked alongside McNally and director Leonard Foglia to create the role at the Ojai Playwrights Conference.[18] He played the lawyer Brad Morris on several seasons of the CBS comedy How I Met Your Mother.[20] In 2008, he joined the cast of The CW drama One Tree Hill for its fifth season, playing bartender Owen Morello. He shot the series in Wilmington, North Carolina, and returned for its sixth and seventh seasons.[21]
Manganiello played Stu on the Fox sitcom 'Til Death for two episodes, and starred in the short film Wounded that year,[22] which he originated on the stage and won Best Short Film at the 2011 Big Island Film Festival.[citation needed] He starred in the direct-to-video war film Behind Enemy Lines: Colombia in 2009 playing Lt. Sean Macklin, a Navy SEAL squad leader. In order to add authenticity to the production, he trained for several months with a former Navy SEAL, for whom he paid to come to the set and stay in the cast's hotel. The film was shot in Puerto Rico.[23] He appeared in an episode of Medium in 2009, and has guest starred on all three series of CBS's CSI television franchise (CSI: Crime Scene Investigation, CSI: Miami, and CSI: NY). He had a role in the independent film Irene in Time. In 2010, he appeared in television commercials for Taco Bell, promoting its new product.[10] He shot television pilots for 100 Questions and the Pittsburgh-set sitcom Livin' on a Prayer.[24][25]
In early 2011, Manganiello was asked to screen test for the title role of Superman in Man of Steel, for which he was rumored to be the front runner, but due to scheduling problems with True Blood, he was forced to drop out during the final stage. He told Access Hollywood in an interview: "They wanted me to screen test and they actually asked for my measurements for the suit and everything... their shoot date switched and it would have taken up 11 weeks out of my True Blood schedule. At the end of the day, we couldn't get the schedule to work... so, regrettably, I never got to screen test, I never got to put the suit on."[28]
Upon completion of filming for season four of True Blood, he shot an episode of USA's White Collar with his former drama school classmate Matt Bomer,[29] and then shot the film adaptation of the best selling book What To Expect When You're Expecting[30] before returning to Los Angeles to shoot an episode of Two and a Half Men with Ashton Kutcher.[31]
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