Janney was born in Dayton, Ohio, the daughter of Macy (née Putnam), a former actress and homemaker, and Jervis Spencer Janney, Jr., a real estate developer and jazz musician. She has one older brother, Jay, and a younger brother, Hal. She attended The Miami Valley School in Dayton, where she got named a distinguished alumna in 2004, and the Hotchkiss School in Connecticut. She then attended Kenyon College. At Kenyon, Janney studied under another Kenyon alum, Paul Newman. She went on to study drama at the Neighborhood Playhouse in New York and the Royal Academy of Dramatic Art in London. She attended The Neighborhood Playhouse the same years as actor Dylan McDermott.
Janney's first roles on television were as comedic foils on soap operas: she first played Vi Kaminski for a short time on As the World Turns, following up with a two-year stint as one of the Spaulding maids, Ginger on Guiding Light. In the spring of 1994, she appeared in the season finale of Law & Order, as a reluctant witness against a member of the Russian mob.
Janney has appeared in a number of films with roles of various sizes, including the 1990s films American Beauty, The Object of My Affection, Big Night, The Impostors, Drop Dead Gorgeous, The Ice Storm, Primary Colors, 10 Things I Hate About You, and Private Parts, and the 2000s films Nurse Betty, The Hours, The Chumscrubber, How to Deal, Winter Solstice and a large role in the animated movie, Finding Nemo, voicing Peach, the starfish. In 2006, Janney carried notable roles in the movies Margaret and Over the Hedge, an animated comedy.
In 1999, she was cast in the role of presidential press secretary C.J. Cregg on the television drama The West Wing, for which she eventually won four Emmy Awards. Two of the Emmys were for Primetime Emmy Award for Outstanding Supporting Actress - Drama Series in 2000 and 2001, and two were for Primetime Emmy Award for Outstanding Lead Actress - Drama Series in the years 2002 and 2004. She was also nominated for Outstanding Lead Actress in a Drama Series in the 2003 and 2006 Primetime Emmys. Janney also won two Screen Actor's Guild awards for Best Actress in a dramatic series, in 2001 and 2002, for her portrayal of C.J Cregg. The cast of The West Wing won the Screen Actor's Guild award for Best Ensemble in a dramatic series the same two years. Additional accolades for Janney's work in The West Wing include four Golden Globe award nominations, and a nomination in 2002 for American Film Institute's Actor of the Year.
Weekend trips to Washington, D.C. were often a part of Janney's schedule, as many outdoor scenes on The West Wing were filmed in the nation's capital. Janney also met several times with former White House Press Secretary Dee Dee Myers who served in the Clinton Administration from 1993–1994. Janney met with Myers in New York City to help portray her character more authentically.
The West Wing concluded in January 2006, and the last episode aired in May of that year. Even though the latter seasons were hit by declining ratings, the overall rating of Janney remained a relatively positive one, touted by Entertainment Weekly as "one to watch", "uncommonly beautiful and infinitely expressive". In January 2006, West Wing's cast was also nominated by the Screen Actors Guild for Outstanding Performance by an Ensemble Cast. Janney, Martin Sheen, Bradley Whitford, Janel Moloney, and other members of the cast appeared at the SAG Awards to honor their late cast mate and friend, John Spencer.
In 2006, she got nominated for an Independent Spirit Award for her performance in the film Our Very Own. Many of Janney's long time friends got involved with Our Very Own, including its producer Shannon McMahon Lichte and cast member Allison Mackie. All three were in the same class at the Neighborhood Playhouse. The writer/director Cameron Watson, also a longtime friend, wrote the role of Joan for her.
In 2007, Janney appeared in the Academy-Award-nominated film, Juno, playing the part of Bren MacGuff, the title character's stepmother, for which she won Best Supporting Actress in the Austin Film Critics Association Awards 2007. In the same year, Janney appeared in the Golden-Globe-nominated film, Hairspray, as Prudy Pingleton, Penny's (Amanda Bynes) strict and religious mother.
Janney appeared in Studio 60 on the Sunset Strip, also created by Aaron Sorkin. She made a guest appearance as herself in The Disaster Show, as the guest host of the week's episode of Studio 60. Many of the characters made references to her part in The West Wing, even those played by fellow West Wing actors.
Janney has remained active in theater. In 1998, she got nominated for a Tony Award for her role in Arthur Miller's A View from the Bridge. Her first Broadway role was in Present Laughter opposite Frank Langella. In 2007, she participated in a workshop for a new musical of the film 9 to 5. In late 2008, Janney joined Broadway stars Stephanie J. Block, Megan Hilty and Marc Kudisch in the new musical, 9 to 5. Based on the film of the same name, Janney is starring as Violet Newstead, the super efficient office manager played by Lily Tomlin in the original film. Janney has signed on for a one-year contract with the 9 to 5 production. She was also nominated for a Tony Award for Best Leading Actress in a Musical for her work in 9 to 5. In May 2009, she won the Drama Desk Award for Outstanding Actress in a Musical for the role of Violet Newstead. She also appeared along Jack Black and John C. Reilly in Prop 8 (The Musical), a satirical musical protesting the passing of California Proposition 8.
In 2004, Janney began lending her voice to television and radio spots created by Kaiser Permanente in the health maintenance organization's broad "Thrive" media campaign, and in a radio campaign for the American Institute of Architects.
In 2010, she appeared as Allison Pearson in In Plain Sight.
In September 2010, it got announced that Janney would be the voice of the Aly San San spokesdroid in the Disney attraction, Star Tours: The Adventure Continues. The attraction later opened at Disney's Hollywood Studios and Disneyland. In the fall of 2013, she became the co-star of the new CBS sitcom, Mom, portraying Bonnie alongside Anna Farris, for which she won her fifth Emmy for Outstanding Performance by an Supporting Actress in a Comedy.
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