Publications by categories in reversed chronological order. * represents co-first authors, ** represents co-corresponding authors.


  1. Allele Sails: launching traits and fates into wild populations with Mendelian DNA sequence modifiers
    Johnson, Michelle L., Hay, Bruce A., and Maselko, Maciej
    bioRxiv 2024
    Hay, Bruce A., Oberhofer, Georg, and Ivy, Tobin
    United States Issued Patent 2024


  1. Cleave and Rescue gamete killers create conditions for gene drive in plants - currently in press at Nature Plants 2024
    Oberhofer, Georg, Johnson, Michelle L., Ivy, Tobin, and Hay, Bruce A.
    bioRxiv 2023
  2. Manipulating the Destiny of Wild Populations Using CRISPR
    Raban, Robyn, Marshall, John, Hay, Bruce A., and Akbari, Omar S.
    Annual Review of Genetics 2023


  1. Gene drive-mediated population elimination for biodiversity conservation. When you come to a fork in the road, take it.
    Hay, Bruce A., and Guo, Ming
    PNAS 2022
  2. Clueless/CLUH regulates mitochondrial fission by promoting recruitment of Drp1 to mitochondria.
    Yang, Huan, Sibilla, Carolina, Liu, Raymond, Yun, Jina, Hay, Bruce A., Blackstone, Craig, Chan, David, Harvey, Robert J, and Guo, Ming
    nature COMMUNICATIONS, 2022


  1. Gene drive that results in addiction to a temperature-sensitive version of an essential gene triggers population collapse in Drosophila.
    Oberhofer, Georg, Ivy, Tobin, and Hay, Bruce A.
    PNAS, 2021
  2. Split versions of Cleave and Rescue selfish genetic elements for measured self limiting gene drive.
    Oberhofer, Georg, Ivy, Tobin, and Hay, Bruce A.
    PLOS Genetics, 2021
  3. Guidelines for the use and interpretation of assays for monitoring autophagy (4th edition).
    Hay, Bruce A., and many, many more
    Autophagy, 2021


  1. Engineering the Composition and Fate of Wild Populations with Gene Drive.
    Hay, Bruce A., Oberhofer, Georg, and Guo, Ming
    Annual Review of Entomology, 2020
    Hay, Bruce A.
    United States Patent Application Publication, 2020
    Hay, Bruce A., and Li, Juan
    United States Patent, 2020
  4. Generation, Analyzing and in-vivo Drug Treatment of Drosophila Models with IBMPFD.
    Zhang, Ting, Hay, Bruce A., and Guo, Ming
    bio-protocol, 2020
  5. A drug-inducible sex-separation technique for insects.
    Kandul, Nikolay P., Hsu, Alexander D., Hay, Bruce A., and Akbari, Omar S.
    Nature Communications, 2020
  6. Gene drive and resilience through renewal with next generation Cleave and Rescue selfish genetic elements.
    Oberhofer, Georg, Ivy, Tobin, and Hay, Bruce A.
    Proceedings of the National Academy of Sciences of the United States of America, 2020


  1. Pest management by genetic addiction.
    Gould, Fred, Dhole, Sumit, and Lloyd, Alun L.
    Proceedings of the National Academy of Sciences of the United States of America, 2019
  2. Cleave and Rescue, a novel selfish genetic element and general strategy for gene drive.
    Oberhofer, Georg, Ivy, Tobin, and Hay, Bruce A.
    Proceedings of the National Academy of Sciences of the United States of America, 2019


    Hay, Bruce A., Oberhofer, Georg, and Ivy, Tobin
    United States Patent Application Publication 2018
  2. Behavior of homing endonuclease gene drives targeting genes required for viability or female fertility with multiplexed guide RNAs.
    Oberhofer, Georg, Ivy, Tobin, and Hay, Bruce A.
    Proceedings of the National Academy of Sciences of the United States of America, 2018
  3. Engineered Reciprocal Chromosome Translocations Drive High Threshold, Reversible Population Replacement in Drosophila.
    Buchman, Anna B., Ivy, Tobin, Marshall, John M., Akbari, Omar S., and Hay, Bruce A.
    ACS Synthetic Biology, 2018
  4. Vectored gene delivery for lifetime animal contraception: Overview and hurdles to implementation.
    Hay, Bruce A., Li, Juan, and Guo, Ming
    Theriogenology, 2018


  1. Rules of the road for insect gene drive research and testing.
    Adelman, et al., and Hay, Bruce A.
    Nature Biotechnology, 2017
  2. A positive readout single transcript reporter for site-specific mRNA cleavage.
    Kandul, Nikolay, Guo, Ming, and Hay, Bruce A.
    PeerJ, 2017
  3. Valosin-containing protein (VCP/p97) inhibitors relieve Mitofusin-dependent mitochondrial defects due to VCP disease mutants.
    Zhang, Ting, Mishra, Prashant, Hay, Bruce A., Chan, David, and Guo, Ming
    eLife, 2017


  1. Selective removal of deletion-bearing mitochondrial DNA in heteroplasmic Drosophila.
    Kandul, Nikolay P., Zhang, Ting, Hay**, Bruce A., and Guo**, Ming
    Nature Communications, 2016
  2. Mapping a multiplexed zoo of mRNA expression.
    Choi, Harry M. T., Calvert, Colby R., Husain, Naeem, Barsi, Julius C., Deverman, Benjamin E., Hunter, Ryan C., Kato, Mihoko, Lee, S. Melanie, Abelin, Anna C. T., Rosenthal, Adam Z., Akbari, Omar S., Li, Yuwei, Hay, Bruce A., Sternberg, Paul W., Patterson, Paul H., Davidson, Eric H., Mazmanian, Sarkis K., Prober, David A., Leadbetter, Jared R., Newman, Dianne K., Readhead, Carol, Bronner, Marianne E., Wold, Barbara, Fraser, Scott E., and Pierce, Niles A.
    Development, 2016


  1. Identification of Genes Uniquely Expressed in the Germ Line Tissues of the Jewel Wasp Nasonia vitripennis.
    Ferree, Patrick M., Fang, Christopher, Mastrodimos, Mariah, Hay, Bruce A., Amrhein, Henry, and Akbari, Omar S.
    G3, 2015
  2. Vectored antibody gene delivery mediates long-term contraception.
    Li, Juan, Olvera, Alejandra I., Akbari, Omar S., Moradian, Annie, Sweredoski, Michael J., Hess, Sonja, and Hay, Bruce A.
    Current Biology, 2015


  1. Novel Synthetic Medea Selfish Genetic Elements Drive Population Replacement in Drosophila; a Theoretical Exploration of Medea-Dependent Population Suppression.
    Akbari, Omar S., Chen, Chun-Hong, Marshall, John M., Huang, Haixia, Antoshechkin, Igor, and Hay, Bruce A.
    ACS Synthetic Biology, 2014
  2. Medusa: A Novel Gene Drive System for Confined Suppression of Insect Populations.
    Marshall, John M., and Hay, Bruce A.
    PLoS ONE, 2014
  3. Identification of germline transcriptional regulatory elements in Aedes aegypti.
    Akbari, Omar S., Papathanos, Philippos A., Sandler, Jeremy E., Kennedy, Katie, and Hay, Bruce A.
    Scientific Reports, 2014


  1. Engineering synthetic medea-based and threshold-dependent underdominance-based gene drive systems in mosquitos.
    Akbari, Omar, Marshall, John, Antoshechkin, Igor, Matzen, Kelly, Papathanos, Philippos, Kennedy, Katie, Ward, Catherine, Cheng, Chun-Hong, Guo, Ming, and Hay, Bruce A.
    Pathogens and Global Health, 2013
  2. Mathematical insights into novel vector control strategies: from outdoor control to confineable transgenic systems.
    Marshall, John, White, Michael, Beier, John, Muller, Gunter, Devine, Gregor, Akbari, Omar, Hay, Bruce, and Ghani, Azra
    Pathogens and Global Health, 2013
  3. A Toolbox of Aedes Aegypti Germline Regulatory Elements for the Development of Genetic Drive Systems and Population Control.
    Papathanos, Philippos A., Akbari, Omar S., and Hay, Bruce A.
    Pathogens and Global Health, 2013
  4. The Developmental Transcriptome of the Mosquito Aedes aegypti, an Invasive Species and Major Arbovirus Vector.
    Akbari, Omar S., Amrhein, Henry, Williams, Brian, Diloreto, Race, Sandler, Jeremy, and Hay, Bruce A.
    G3, 2013
  5. Transcriptome Profiling of Nasonia vitripennis Testis Reveals Novel Transcripts Expressed from the Selfish B Chromosome, Paternal Sex Ratio.
    Akbari, Omar S., Antoshechkin, Igor, Hay, Bruce A., and Ferree, Patrick M.
    G3, 2013
  6. A Synthetic Gene Drive System for Local, Reversible Modification and Suppression of Insect Populations.
    Akbari, Omar S., Matzen, Kelly D., Marshall, John M., Huang, Haixa, Ward, Catherine M., and Hay, Bruce A.
    Current Biology, 2013
  7. Essential role of grim-led programmed cell death for the establishment of corazonin-producing peptidergic nervous system during embryogenesis and metamorphosis in Drosophila melanogaster.
    Lee, Gyunghee, Sehgal, Ritika, Wang, Zixing, Nair, Sudershana, Kikuno, Keiko, Chen, Chun-Hong, Hay, Bruce, and Park, Jae H.
    Biology Open, 2013


  1. General principles of single-construct chromosomal gene drive.
    Marshall, John M., and Hay, Bruce A.
    Evolution, 2012
  2. Confinement of gene drive systems to local populations: A comparative analysis.
    Marshall, John M., and Hay, Bruce A.
    Journal of Theoretical Biology, 2012
  3. Molecular Insights into Parkinson’s Disease. In: Molecular Biology of Neurodegenerative Diseases.
    Rochet, Jean-Christophe, Hay, Bruce A., and Guo, Ming
    Progress in Molecular Biology and Translational Science, 2012


  1. Inverse Medea as a Novel Gene Drive System for Local Population Replacement: A Theoretical Analysis.
    Marshall, John M., and Hay, Bruce A.
    Journal of Heredity, 2011
  2. Medea Selfish Genetic Elements as Tools for Altering Traits of Wild Populations: A Theoretical Analysis.
    Ward, Catherine M., Su, Jessica T., Huang, Yunxin, Lloyd, Alun L., Gould, Fred, and Hay, Bruce A.
    Evolution, 2011
  3. Semele: A Killer-Male, Rescue-Female System for Suppression and Replacement of Insect Disease Vector Populations.
    Marshall, John M., Pittman, Geoffrey W., Buchman, Anna B., and Hay, Bruce A.
    Genetics, 2011
  4. Drosophila caspases involved in developmentally regulated programmed cell death of peptidergic neurons during early metamorphosis.
    J Comp Neurol., 2011


  1. Engineering the genomes of wild insect populations: Challenges, and opportunities provided by synthetic Medea selfish genetic elements.
    Hay, Bruce A., Chen, Chun-Hong, Ward, Catherine M., Huang, Haixia, Su, Jessica T., and Guo, Ming
    Journal of Insect Physiology, 2010
  2. Inactivation of Both foxo and reaper Promotes Long-Term Adult Neurogenesis in Drosophila.
    Siegrist, Sarah E., Haque, Najm S., Chen, Chun-Hong, Hay, Bruce A., and Hariharan, Iswar K.
    Current Biology, 2010


  1. The deubiquitinase emperor’s thumb is a regulator of apoptosis in Drosophila.
    Ribaya, Jeronimo P., Ranmuthu, Madhuka, Copeland, Jeff, Boyarskiy, Sergey, Blair, Adrienne P., Hay, Bruce, and Laski, Frank A.
    Developmental Biology, 2009


  1. Identification of novel genes involved in light-dependent CRY degradation through a genome-wide RNAi screen.
    Genes Dev., 2008
  2. Differential requirements for the Pax6(5a) genes eyegone and twin of eyegone during eye development in Drosophila.
    Developmental Biology, 2008


  1. Stage-specific differences in the requirements for germline stem cell maintenance in the Drosophila ovary.
    Shcherbata, Halyna R., Ward, Ellen J., Fischer, Karin A., Yu, Jenn-Yah, Reynolds, Steven H., Chen, Chun-Hong, Xu, Peizhang, Hay, Bruce A., and Ruohola-Baker, Hannele
    Cell Stem Cell, 2007
  2. echinus, required for interommatidial cell sorting and cell death in the Drosophila pupal retina, encodes a protein with homology to ubiquitin-specific proteases.
    Copeland, Jeffrey M., Bosdet, Ian, Freeman, J. Douglas, Guo, Ming, Gorski, Sharon M., and Hay, Bruce A.
    BMC Developmental Biology, 2007
  3. A Synthetic Maternal-Effect Selfish Genetic Element Drives Population Replacement in Drosophila.
    Chen, Chun-Hong, Huang, Haixia, Ward, Catherine M., Su, Jessica T., Schaeffer, Lorian V., Guo, Ming, and Hay, Bruce A.
    Science, 2007
  4. The Drosophila Inhibitor of Apoptosis (IAP) DIAP2 Is Dispensable for Cell Survival, Required for the Innate Immune Response to Gram-negative Bacterial Infection, and Can Be Negatively Regulated by the Reaper/Hid/Grim Family of IAP-binding Apoptosis Inducers.
    Huh, Jun R., Foe, Ian, Muro, Israel, Chen, Chun Hong, Seol, Jae Hong, Yoo, Soon Ji, Guo, Ming, Park, Jin Mo, and Hay, Bruce A.
    Journal of Biological Chemistry, 2007


  1. Drosophila caspases involved in developmentally regulated programmed cell death of peptidergic neurons during early metamorphosis.
    Chen, Chun-Hong, Guo, Ming, and Hay, Bruce A.
    Humana Press, 2006
  2. Caspase-Dependent Cell Death in Drosophila.
    Hay, Bruce A., and Guo, Ming
    Annual Review of Cell and Developmental Biology, 2006
  3. The Drosophila caspase Ice is important for many apoptotic cell deaths and for spermatid individualization, a nonapoptotic process.
    Muro, Israel, Berry, Deborah L., Huh, Jun R., Chen, Chun Hong, Huang, Haixia, Yoo, Soon Ji, Guo, Ming, Baehrecke, Eric H., and Hay, Bruce A.
    Development, 2006
  4. Drosophila pink1 is required for mitochondrial function and interacts genetically with parkin.
    Clark, Ira E., Dodson, Mark W., Cao, Joseph H., Huh, Jun R., Seol, Jae Hong, Yoo, Soon Ji, Hay, Bruce A., and Guo, Ming
    Nature, 2006
  5. Structure and Activation Mechanism of the Drosophila Initiator Caspase Dronc.
    Yan, Nieng, Huh, Jun R., Schirf, Virgil, Demeler, Borries, Hay, Bruce A., and Shi, Yigong
    Journal of Biological Chemistry, 2006


  1. The genetics of cell death: approaches, insights and opportunities in Drosophila.
    Hay, Bruce A., Huh, Jun R., and Guo, Ming
    Nature Reviews. Genetics, 2004
  2. MicroRNAs and the regulation of cell death.
    Xu, Peizhang, Guo, Ming, and Hay, Bruce A.
    TRENDS in Genetics, 2004
  3. Compensatory Proliferation Induced by Cell Death in the Drosophila Wing Disc Requires Activity of the Apical Cell Death Caspase Dronc in a Nonapoptotic Role.
    Huh, Jun R., Guo, Ming, and Hay, Bruce A.
    Current Biology, 2004
  4. Multiple Apoptotic Caspase Cascades Are Required in Nonapoptotic Roles for Drosophila Spermatid Individualization.
    Huh, Jun R., Vernooy, Stephanie Y., Yu, Hong, Yan, Nieng, Shi, Yigong, Guo, Ming, and Hay, Bruce A.
    PLoS Biology, 2004


  1. Molecular mechanism of Reaper-Grim-Hid-mediated suppression of DIAP1-dependent Dronc ubiquitination.
    Chai, Jijie, Yan, Nieng, Huh, Jun R., Wu, Jia-Wei, Li, Wenyu, Hay, Bruce A., and Shi, Yigong
    Nature Structural Biology, 2003
  2. Coupling Cell Growth, Proliferation, and Death: Hippo Weighs In.
    Guo, Ming, and Hay, Bruce A.
    Developmentl Cell, 2003
  3. A reporter for amyloid precursor protein γ-secretase activity in Drosophila.
    Guo, Ming, Hong, Elizabeth J., Fernandes, Jolene, Zipursky, S. Larry, and Hay, Bruce A.
    Human Molecular Genetics, 2003
  4. The Drosophila MicroRNA Mir-14 Suppresses Cell Death and Is Required for Normal Fat Metabolism.
    Xu, Peizhang, Vernooy, Stephanie Y., Guo, Ming, and Hay, Bruce A.
    Current Biology, 2003
  5. Reaper is regulated by IAP-mediated ubiquitination.
    Olson, Michael R., Holley, Christopher L., Yoo, Soon Ji, Huh, Jun R., Hay, Bruce A., and Kornbluth, Sally
    Journal of Biological Chemistry, 2003


  1. The Drosophila DIAP1 protein is required to prevent accumulation of a continuously generated, processed form of the apical caspase DRONC.
    Muro, Israel, Hay, Bruce A., and Clem, Rollie J.
    Journal of Biological Chemistry, 2002
  2. Apoptosis: sculpture of a fly’s head.
    Huh, Jun R., and Hay, Bruce A.
    Nature News and Views, 2002
  3. Drosophila Bruce can potently suppress Rpr- and Grim-dependent but not Hid-dependent cell death.
    Vernooy, Stephanie Y., Chow, Vivian, Su, Julius, Verbrugghe, Koen, Yang, Jennifer, Cole, Susannah, Olson, Michael R., and Hay, Bruce A.
    Current Biology, 2002
  4. A pathway of signals regulating effector and initiator caspases in the developing Drosophila eye.
    Yu, Sun-Yun, Yoo, Soon Ji, Yang, Lihui, Zapata, Cynthia, Srinivasan, Anu, Hay, Bruce A., and Baker, Nicholas E.
    Development, 2002
  5. Hid, Rpr and Grim negatively regulate DIAP1 levels through distinct mechanisms.
    Yoo, Soon Ji, Huh, Jun R., Muro, Israel, Yu, Hong, Wang, Lijuan, Wang, Susan L., Feldman, R. M. Renny, Clem, Rollie J., Müller, H.-Arno J., and Hay, Bruce A.
    Nature Cell Biology, 2002
  6. The role of cytochrome c in caspase activation in Drosophila melanogaster cells.
    Dorstyn, Loretta, Read, Stuart, Cakouros, Dimitrios, Huh, Jun R., Hay, Bruce A., and Kumar, Sharad
    Journal of Cell Biology, 2002


  1. Structural analysis of a functional DIAP1 fragment bound to grim and hid peptides.
    Wu, Jia-Wei, Cocina, Amy E., Chai, Jijie, Hay, Bruce A., and Shi, Yigong
    Molecular Cell, 2001


  1. Understanding IAP function and regulation: a view from Drosophila.
    Hay, Bruce A.
    Cell Death and Differentiation, 2000
  2. Monitoring activity of caspases and their regulators in yeast Saccharomyces cerevisiae.
    Hawkins, Christine J., Wang, Susan L., and Hay, Bruce A.
    Methods in Enzymology, 2000
  3. Cell death regulation in Drosophila: Conservation of mechanism and unique insights.
    Vernooy, Stephanie Y., Copeland, Jeffrey, Ghaboosi, Nazli, Griffin, Erik E., Yoo, Soon Ji, and Hay, Bruce A.
    Journal of Cell Biology, 2000
  4. The Drosophila Caspase DRONC Cleaves following Glutamate or Aspartate and Is Regulated by DIAP1, HID, and GRIM.
    Hawkins, Christine J., Yoo, Soon Ji, Peterson, Erin P., Wang, Susan L., Vernooy, Stephanie Y., and Hay, Bruce A.
    Journal of Biological Chemistry, 2000
  5. Comparative Genomics of the Eukaryotes.
    Rubin, et al., and Hay, Bruce A.
    Science, 2000


  1. The Drosophila Caspase Inhibitor DIAP1 Is Essential for Cell Survival and Is Negatively Regulated by HID.
    Wang, Susan L., Hawkins, Christine J., Yoo, Soon Ji, Müller, H.-Arno J., and Hay, Bruce A.
    Cell, 1999
  2. A cloning method to identify caspases and their regulators in yeast: Identification of Drosophila IAP1 as an inhibitor of the Drosophila caspase DCP-1.
    Hawkins, Christine J., Wang, Susan L., and Hay, Bruce A.
    Proceedings of the National Academy of Sciences of the United States of America, 1999


  1. P element insertion-dependent gene activation in the Drosophila eye.
    Hay, Bruce A., Maile, Randy, and Rubin, Gerald M.
    Proceedings of the National Academy of Sciences of the United States of America, 1997


  1. Drosophila Homologs of Baculovirus Inhibitor of Apoptosis Proteins Function to Block Cell Death.
    Hay, Bruce A., Wassarman, David A., and Rubin, Gerald M.
    Cell, 1995


  1. Expression of baculovirus P35 prevents cell death in Drosophila.
    Hay, Bruce A., Wolff, Tanya, and Rubin, Gerald M.
    Development, 1994


  1. The germ cell-less gene product: a posteriorly localized component necessary for germ cell development in Drosophila.
    Jongens, Thomas A., Hay, Bruce A., Jan, Lily Y., and Jan, Yuh N.
    Cell, 1992


  1. Localization of vasa, a component of Drosophila polar granules, in maternal-effect mutants that alter embryonic anteroposterior polarity.
    Hay, Bruce, Jan, Lily Yeh, and Jan, Yuh Nung
    Development, 1990


  1. A protein component of Drosophila polar granules is encoded by vasa and has extensive sequence similarity to ATP-dependent helicases.
    Hay, Bruce, Jan, Lily Yeh, and Jan, Yuh Nung
    Cell, 1988
  2. Identification of a component of Drosophila polar granules.
    Hay, Bruce, Ackerman, Larry, Barbel, Sandra, Jan, Lily Y., and Jan, Yuh Nung
    Development, 1988


  1. Evidence for a secretory form of the cellular prion protein.
    Hay, Bruce A., Prusiner, Stanley B., and Lingappa, Vishwanath R.
    Biochemsitry, 1987
  2. Biogenesis and transmembrane orientation of the cellular isoform of the scrapie prion protein.
    Hay, Bruce A., Barry, Ronald A., Liebergurg, Ivan, Prusiner, Stanley B., and Lingappa, Vishwanath R.
    Mol.Cell.Biol., 1987
  3. Developmental expression of the prion protein in brain.
    McKinley, Michael P., Hay, Bruce A., Lingappa, Vishwanath R., Lieberburg, Ivan, and Prusiner, Stanley B.
    Dev. Biol., 1987


  1. Depolarization-induced effects of Ca2+-calmodulin-dependent protein kinase injection, in vivo, in single neurons of cat motor cortex.
    Woody, Charles D., Alkon, Daniel L., and Hay, Bruce A.
    Brain Res., 1984
  2. Voltage-dependent calcium and calcium-activated potassium currents of a molluscan photoreceptor.
    Alkon, Daniel L., Farley, Joseph, Sakakibara, Manabu, and Hay, Bruce A.
    Biophys. J., 1984