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Originally published as Genetics Published Articles Ahead of Print on August 24, 2007.
Genetics, Vol. 177, 689-697, October 2007, Copyright © 2007
doi:10.1534/genetics.107.077735
Genomewide Clonal Analysis of Lethal Mutations in the Drosophila melanogaster Eye: Comparison of the X Chromosome and Autosomes
Gerald B. Call*,1,2,
John M. Olson*,1,
Jiong Chen*,3,
Nikki Villarasa*,
Kathy T. Ngo*,
Allison M. Yabroff*,4,
Shawn Cokus*,
Matteo Pellegrini*,
Elena Bibikova*,
Chris Bui*,
Albert Cespedes*,
Cheryl Chan*,
Stacy Chan*,
Amrita K. Cheema*,
Akanksha Chhabra*,
Vida Chitsazzadeh*,
Minh-Tu Do*,
Q. Angela Fang*,
Andrew Folick*,
Gelsey L. Goodstein*,
Cheng R. Huang*,
Tony Hung*,
Eunha Kim*,
William Kim*,
Yulee Kim*,
Emil Kohan*,
Edward Kuoy*,
Robert Kwak*,
Eric Lee*,
JiEun Lee*,
Henry Lin*,
H-C. Angela Liu*,
Tatiana Moroz*,
Tharani Prasad*,
Sacha L. Prashad*,
Alexander N. Patananan*,
Alma Rangel*,
Desiree Rosselli*,
Sohrab Sidhu*,
Daniel Sitz*,
Chelsea E. Taber*,
Jingwen Tan*,
Kasey Topp*,
PhuongThao Tran*,
Quynh-Minh Tran*,
Mary Unkovic*,
Maggie Wells*,
Jessica Wickland*,
Kevin Yackle*,
Amir Yavari*,
Jesse M. Zaretsky*,
Christopher M. Allen*,
Latifat Alli*,
Ju An*,
Abbas Anwar*,
Sonia Arevalo*,
Danny Ayoub*,
Shawn S. Badal*,
Armonde Baghdanian*,
Arthur H. Baghdanian*,
Sara A. Baumann*,
Vivian N. Becerra*,
Hei J. Chan*,
Aileen E. Chang*,
Xibin A. Cheng*,
Mabel Chin*,
Fleurette Chong*,
Carlyn Crisostomo*,
Sanjit Datta
,
Angela Delosreyes*,
Francie Diep*,
Preethika Ekanayake*,
Mark Engeln*,
Elizabeth Evers*,
Farzin Farshidi*,
Katrina Fischer*,
Arlene J. Formanes*,
Jun Gong*,
Riju Gupta*,
Blake E. Haas*,
Vicky Hahm*,
Michael Hsieh*,
James Z. Hui*,
Mei L. Iao*,
Sophia D. Jin*,
Angela Y. Kim*,
Lydia S-H. Kim*,
Megan King
,
Chloe Knudsen-Robbins*,
David Kohanchi*,
Bogdana Kovshilovskaya*,
Amy Ku*,
Raymond W. Kung*,
Mark E. L. Landig*,
Stephanie S. Latterman*,
Stephanie S. Lauw*,
Daniel S. Lee*,
Joann S. Lee*,
Kai C. Lei*,
Lesley L. Leung*,
Renata Lerner*,
Jian-ya Lin*,
Kathleen Lin*,
Bryon C. Lim*,
Crystal P. Y. Lui*,
Tiffany Q. Liu*,
Vincent Luong*,
Jacob Makshanoff*,
An-Chi Mei*,
Miguel Meza*,
Yara A. Mikhaeil*,
Majid Moarefi*,
Long H. Nguyen*,
Shekhar S. Pai*,
Manish Pandya*,
Aadit R. Patel*,
Paul D. Picard
,
Michael M. Safaee*,
Carol Salame*,
Christian Sanchez*,
Nina Sanchez*,
Christina C. Seifert*,
Abhishek Shah*,
Oganes H. Shilgevorkyan*,
Inderroop Singh*,
Vanessa Soma*,
Junia J. Song*,
Neetika Srivastava*,
Jennifer L. Sta.Ana*,
Christie Sun*,
Diane Tan*,
Alison S. Teruya*,
Robyn Tikia*,
Trinh Tran*,
Emily G. Travis*,
Jennifer D. Trinh*,
Diane Vo*,
Thomas Walsh*,
Regan S. Wong*,
Katherine Wu*,
Ya-Whey Wu*,
Nkau X. V. Yang*,
Michael Yeranosian*,
James S. Yu*,
Jennifer J. Zhou*,
Ran X. Zhu*,
Anna Abrams*,
Amanda Abramson*,
Latiffe Amado*,
Jenny Anderson*,
Keenan Bashour*,
Elsa Beyer*,
Allen Bookatz*,
Sarah Brewer*,
Natalie Buu*,
Stephanie Calvillo*,
Joseph Cao*,
Amy Chan*,
Jenny Chan*,
Aileen Chang*,
Daniel Chang*,
Yuli Chang,
YiBing Chen*,
Joo Choi*,
Jeyling Chou*,
Peter Dang*,
Sumit Datta*,
Ardy Davarifar*,
Artemis Deravanesian*,
Poonam Desai*,
Jordan Fabrikant*,
Shahbaz Farnad*,
Katherine Fu*,
Eddie Garcia*,
Nick Garrone*,
Srpouhi Gasparyan*,
Phyllis Gayda*,
Sherrylene Go*,
Chad Goffstein*,
Courtney Gonzalez*,
Mariam Guirguis*,
Ryan Hassid*,
Brenda Hermogeno*,
Julie Hong*,
Aria Hong*,
Lindsay Hovestreydt*,
Charles Hu*,
Devon Huff*,
Farid Jamshidian*,
James Jen*,
Katrin Kahen*,
Linda Kao*,
Melissa Kelley*,
Thomas Kho*,
Yein Kim*,
Sarah Kim*,
Brian Kirkpatrick*,
Adam Langenbacher*,
Santino Laxamana*,
Janet Lee*,
Chris Lee*,
So-Youn Lee*,
ToHang S. Lee*,
Toni Lee*,
Gemma Lewis*,
Sheila Lezcano*,
Peter Lin*,
Thanh Luu*,
Julie Luu*,
Will Marrs*,
Erin Marsh*,
Jamie Marshall*,
Sarah Min*,
Tanya Minasian*,
Helena Minye*,
Amit Misra*,
Miles Morimoto*,
Yasaman Moshfegh*,
Jessica Murray*,
Kha Nguyen*,
Cynthia Nguyen*,
Ernesto Nodado, II*,
Amanda O'Donahue*,
Ndidi Onugha*,
Nneka Orjiakor*,
Bhavin Padhiar*,
Eric Paul*,
Mara Pavel-Dinu*,
Alex Pavlenko*,
Edwin Paz*,
Sarah Phaklides*,
Lephong Pham*,
Preethi Poulose*,
Russell Powell*,
Aya Pusic*,
Divi Ramola*,
Kirsten Regalia*,
Meghann Ribbens*,
Bassel Rifai*,
Manyak Saakyan*,
Pamela Saarikoski*,
Miriam Segura*,
Farnaz Shadpour*,
Aram Shemmassian*,
Ramnik Singh*,
Vivek Singh*,
Emily Skinner*,
Daniel Solomin*,
Kosha Soneji*,
Kristin Spivey*,
Erika Stageberg*,
Marina Stavchanskiy*,
Leena Tekchandani*,
Leo Thai*,
Jayantha Thiyanaratnam*,
Maurine Tong*,
Aneet Toor*,
Steve Tovar*,
Kelly Trangsrud*,
Wah-Yung Tsang*,
Marc Uemura*,
Emily Vollmer*,
Emily Weiss*,
Damien Wood*,
Joy Wu*,
Sophia Wu*,
Winston Wu*,
Qing Xu*,
Yuki Yamauchi*,
Will Yarosh*,
Laura Yee*,
George Yen* and
Utpal Banerjee*,**,
,5
* Department of Molecular, Cell, and Developmental Biology, University of California, Los Angeles, California 90095,
Palos Verdes Peninsula High School, Rolling Hills Estates, California 90274,
Culver City Independent Study School, Culver City, California 90230,
Loyola High School, Los Angeles, California 90006, ** Molecular Biology Institute, University of California, Los Angeles, California 90095 and 
Department of Biological Chemistry, University of California, Los Angeles, California 90095
5 Corresponding author: Department of Molecular, Cell, and Developmental Biology, 2204 Life Science, 621 Charles E. Young Dr. South, Los Angeles, CA 90095.
E-mail: banerjee{at}mbi.ucla.edu
Using a large consortium of undergraduate students in an organized program at the University of California, Los Angeles (UCLA), we have undertaken a functional genomic screen in the Drosophila eye. In addition to the educational value of discovery-based learning, this article presents the first comprehensive genomewide analysis of essential genes involved in eye development. The data reveal the surprising result that the X chromosome has almost twice the frequency of essential genes involved in eye development as that found on the autosomes.
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