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THE HELEN HAY WHITNEY FOUNDATION

2014-2015 Annual Report

20 Squadron Boulevard, Suite 630 New City, NY 10956 www.hhwf.org Tel: (845) 639-6799 Fax: (845) 639-6798 THE HELEN HAY WHITNEY FOUNDATION

BOARD OF TRUSTEES

Averil Payson Meyer, President Steven C. Harrison, Ph.D., Vice President Lisa A. Steiner, M.D., Vice President W. Perry Welch, Treasurer Thomas M. Jessell, Ph.D. Payne W. Middleton Thomas P. Sakmar, M.D. Stephen C. Sherrill

SCIENTIFIC ADVISORY COMMITTEE

Steven C. Harrison, Ph.D., Chairman David J. Anderson, Ph.D. Daniel Kahne, Ph.D. Philippa Marrack, Ph.D. Markus Meister, Ph.D. Barbara J. Meyer, Ph.D. Julie A. Theriot, Ph.D. Jonathan S. Weissman, Ph.D. S. Lawrence Zipursky, Ph.D.

ADMINISTRATIVE DIRECTOR and SECRETARY

Robert Weinberger

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REPORT OF THE VICE PRESIDENT AND CHAIRMAN, SCIENTIFIC ADVISORY COMMITTEE

I am pleased to report on two years of activity for the Scientific Advisory Committee. Our charge, to select an outstanding group of Fellows each year and to respond as needed to advisory issues that occasionally arise during the tenure of their fellowship, is always a stimulating one, and the talks from third-year Fellows at the Annual Meeting continue to give remarkably positive feedback.

Our partnerships with HHMI, the Simons Foundation, and Merck have benefitted the program greatly. We have three Simons Fellows, eight HHMI Fellows and two Merck Fellows, in addition to the eleven we now support from endowment income. (Let me add, on behalf of the SAC, a note of thanks to Perry Welch, the Treasurer of the Foundation, who oversees that endowment with great dedication and insight.) A class of 24 means the competition is fierce, and we continue to identify an extraordinary group of young biomedical scientists.

Membership in the SAC has not changed since Jonathan Weissman, Markus Meister, and Pippa Marrack came on board, as I reported in my last SAC Chair's report. We have a gratifyingly dedicated, lively, and deeply thoughtful team. Our annual deliberations become lively discussions of a splendidly broad range of science, while never losing sight of the critical dimensions of talent and promise we look for in our candidates. I hope we continue to choose as wisely as did our predecessors: Cori Bargmann (HHWF '90) and Bob Weinberg (HHWF '72) were recipients of the new in Life Sciences in 2013, Tom Lindahl (HHWF '70) shared the 2015 Nobel Prize in Chemistry, and Larry Zipursky (HHWF'87), and a current member of our SAC, received the 2015 Louisa Gross Horwitz Prize.

As I look at the state of biomedical research in the US, I do worry that it will be harder for the current class of Whitney Fellows to thrive as easily as did the generation that built careers at a time of reasonable consensus about the role of science in our society and the importance of governmental support for its pursuit. I am confident that the Fellows we select will be the survivors in a context far more challenging that the one faced by Fellows who started their independent careers in the '80's and 90's.

The SAC would not be able to work as effectively in selecting such outstanding Fellows without Robert Weinberger, whose deft administrative skills and thoughtful attention to the personal needs of our Fellows. His management of our operation makes our success possible.

Stephen C. Harrison, Ph.D.

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ORGANIZATION

The Foundation was established and endowed by Mrs. Charles S. Payson, the former Joan Whitney, in 1947, and named in honor of her mother, Helen Hay Whitney. The current Postdoctoral Research Fellowship Program was begun in 1956.

The Foundation is a New York State not-for-profit corporation. The corporation elects a Board of Trustees of not more than eleven members at its annual meeting. Full power in the management and control of the affairs of the corporation is vested in this Board, which holds two stated meetings a year. An Executive Committee acts for the Board in the intervals between meetings.

The Postdoctoral Research Fellowship Program is the responsibility of the Scientific Advisory Committee and is under its continual review and evaluation. The day-to-day administration of the fellowship program is the responsibility of the Administrative Director, who reports to the Chairman of the Scientific Advisory Committee.

PURPOSE

The Foundation was established to stimulate and support research in the area of rheumatic fever and rheumatic heart disease, later expanding its interests to include diseases of connective tissue and, ultimately, all basic biomedical sciences. Under the Foundation’s program, young men and women planning careers in biological or medical research receive financial support of sufficient duration to help further their professional careers. Residents of the , Canada, and Mexico who hold the M.D., Ph.D., or equivalent degree, or US citizens residing abroad, are eligible to apply for a fellowship for beginning postdoctoral training in biomedical research.

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THE WALTER BAUER MEMORIAL MEETING

The Annual meeting of Fellows is held in November each year, and at this meeting each third-year Fellow speaks on the results of his or her research during the tenure of the fellowship. The Foundation also invites a distinguished scientist to present the Maclyn McCarty Annual Lecture.

The Fifty-Seventh Annual Fellows Meeting was held at the MIT Endicott House, November 7-9, 2014, with 58 current Fellows in attendance as well as the Scientific Advisory Committee, members of the Board of Trustees, and the Administrative Director. There were 17 presentations by the Fellows in their final year of fellowship support. Hidde Ploegh, Ph.D. Member of the Whitehead Institute, and Professor of Biology at the Massachusetts Institute of Technology, presented the Maclyn McCarty lecture.

The Fifty-Eighth Annual Fellows Meeting was held at the MIT Endicott House, November 6-8, 2015, with 63 current Fellows in attendance as well as the Scientific Advisory Committee, members of the Board of Trustees, and the Administrative Director. There were 20 presentations by the Fellows in their final year of fellowship support. David Julius, Ph.D. Professor and Chair, at the Department of , University of , San Francisco, presented the Maclyn McCarty lecture.

The Foundation continues to think that the annual meeting is an important component of the fellowship program because of the opportunity it provides our Fellows for scientific interchange and exposure to biomedical fields other than their own.

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The Helen Hay Whitney Foundation Final - Year Fellows Fifty-Seventh Annual Fellows Meeting - November 7-9, 2014

Top row, left to right: Eliezer Calo-Velazquez, Richard K. Hite, Alicia A. Bicknell, Andres M. Lebensohn, Sergei Doulatov, Paula D. Montero Llopis, Nathan W. Pierce, Andrés Bendesky, Nuo Li.

Front row, left to right: Ying Liu, Eleni Mimitou, Richard N. McLaughlin, Saori L. Haigo, Rebecca L. Lamason, Megan L. Matthews, Ming Wu.

Absent from photo: Dmitriy Aronov

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The Helen Hay Whitney Foundation Final - Year Fellows Fifty-Eighth Annual Fellows Meeting - November 6-8, 2015

Top row, left to right: Anna B. Loveland, Andrew Miri, Cristian Boboila, Zachary B. Hill, Michel J. DuPage, Michael B. Lazarus.

Front row, left to right: Stephen N. Floor, Elena Piskounova, Vidhya Ramachandran, Saher Sue Hammoud, Daniel J. Dickinson, Aaron S. Andalman, Weizhe Hong.

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LOCATION OF FELLOWS COMPLETING FELLOWSHIP TENURE

Those who completed their fellowships during calendar year 2014, are listed below, together with their post-fellowship locations.

Eiman Azim, Ph.D. Assistant Professor, Molecular Neurobiology Laboratory, Salk Institute for Biological Studies.

Maya Bader, Ph.D. Scientific Grants Manager at the Pancreatic Cancer Action Network, .

Joshua J. Bayes, Ph.D. Product Manager at Adaptive Biotechnologies Corp., San Franciso.

Alicia A. Bicknell, Ph.D. Postdoctoral Fellow at University of Massachusetts Medical School

Stephen G. Brohawn, Ph.D. Assistant Professor of Neurobiology, Department of Molecular and Cell Biology & Helen Wills Institute, University of California, Berkeley.

Sergei Doulatov, Ph.D. Assistant Professor of , Division of Hematology, University of Washington School of Medicine.

Evan H. Feinberg, Ph.D. Assistant Professor, Department of Anatomy, University of California, San Francisco.

Walter M. Fischler, Ph.D. Postdoctoral Fellow, Axel Lab, .

Lillian K. Fritz-Laylin, Ph.D. Postdoctoral Fellow, Mullins Lab, University of California, San Francisco.

Saher Sue Hammoud, Ph.D. Assistant Professor of Human Genetics, Assistant Professor of Urology, University of Michigan.

Karen E. Kasza, Ph.D. Assistant Professor, Department of Mechanical Engineering, Columbia University.

Andres M. Lebensohn, Ph.D. Postdoctoral Fellow, Rohatgi Lab, Stanford University.

Ying Liu, Ph.D. Principle Investigator, Institute of Molecular Medicine, Peking University, Beijing, China.

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Colin D. Malone, Ph.D. Director of Genomic Analysis and Technical Operations as the Institute for Genomic Medicine, Assistant Professor of Medical Sciences, Columbia University.

Michael B. Manookin, Ph.D. Acting Instructor, University of Washington, Department of Ophthalmology, Seattle, WA

Lisa F. Marshall, Ph.D. Senior Research Scientist at Daktari Diagnostics, Cambridge, MA.

Yasemin S. Sancak, Ph.D. Postdoctoral Fellow, Mootha Lab, Massachusetts General Hospital.

Xin Ye, Ph.D. Postdoctoral Fellow, Weinberg Lab, Whitehead Institute for Biomedical Research.

Xin Zhang, Ph.D. Assistant Professor of Chemistry, and Molecular Biology, The Pennsylvania State University.

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LOCATION OF FELLOWS COMPLETING FELLOWSHIP TENURE

Those who completed their fellowships during calendar year 2015, are listed below, together with their post-fellowship locations.

Aaron S. Andalman, Ph.D. Chief Science Officer and Co-Founder, Cognitiv, New York, NY.

Dmitriy Aronov, Ph.D. Postdoctoral Fellow, Tank Lab, .

Andrés Bendesky, M.D., Ph.D. Principal Investigator, Zuckerman Mind Brain Behavior Institute, Columbia University.

Michael E. Birnbaum, Ph.D. Assistant Professor of Biological Engineering, Koch Institute for Integrative Cancer Research at MIT.

Eliezer Calo-Velazquez, Ph.D. Postdoctoral Fellow, Wysocka Lab, Stanford University School of Medicine.

Margaret F. Carr-Larkin, Ph.D. Consultant, The Boston Consulting Group, Washington, DC.

Daniel J. Dickinson, Ph.D. Postdoctoral Fellow, Goldstein lab, The University of North Carolina at Chapel Hill.

Christopher B. Ford, Ph.D. Senior Scientist, Microbiome Sciences at Seres Therapeutics, Cambridge, MA.

Saori L. Haigo, Ph.D. Physician Partner at UCLA Health.

Richard K. Hite Jr., Ph.D. Principal Investigator, Structural Biology Program, Sloan Kettering Institute

Rebecca L. Lamason, Ph.D. Postdoctoral Fellow, Welch Lab, University of California, Berkeley.

Lishi Li, Ph.D. Junior Associate at McKinsey & Company, Shanghai City, China.

Nuo Li, Ph.D. Assistant Professor, Neuroscience, Baylor College of Medicine.

Megan L. Matthews, Ph.D. Postdoctoral Fellow, Cravatt lab, The Scripps Research Institute.

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Richard N. McLaughlin, Ph.D. Postdoctoral Fellow, Malik lab, Fred Hutchinson Cancer Research Center.

Eleni Mimitou, Ph.D. Postdoctoral Fellow, Keeney lab, Memorial Sloan-Kettering Cancer Center.

Paula D. Montero Llopis, Ph.D. Microscopy Research Associate/ MicRoN manager, Harvard Medical School.

Nathan W. Pierce, Ph.D. Scientist at Verily (Google Life Sciences) Mountain View, CA.

Cindy Poo, Ph.D. Postdoctoral Fellow, Mainen lab, Champalimaud Center for the Unknown, Portugal.

Ming Wu, Ph.D. Senior Scientist, Salubris Biotherapeutics, Inc., Gaithersburg, MD.

Xin Xiong, Ph.D. Research Scientist, Agenovir, San Francisco, CA.

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LIST OF AWARDS FOR 2014

Of the 471 applications received in 2014, 459 (99%) were from candidates with the Ph.D. or comparable degree awarded or pending, 7 from candidates with the M.D./Ph.D. degree, and 3 were from candidates with the M.D. degree. Of these applicants, 41 qualified for personal interview, and 25 were awarded fellowships.

Michael E. Birnbaum, Ph.D. For training at Stanford University with Dr. Carla Shatz. Characterization and manipulation of the PirB-ABeta interaction, a novel pathway in Alzheimer’s Disease.

Emily Jordan Capra, Ph.D. For training at the California Institute of Technology with Dr. Michael Elowitz. Notch signaling from the bottom up: mapping directionality and specificity of intercellular signaling.

Colin C. Conine, Ph.D. For training at the University of Massachusetts Medical School - Worcester with Dr. Oliver Rando. The role of paternally-transmitted small RNAs in reprogramming preimplantation development.

Rachel A. Duffié, Ph.D. For training at Columbia University with Dr. Stavros Lomvardas. Regulatory Noncoding RNAs in the Olfactory Sensory Epithelium.

Ellen R. Edenberg, Ph.D. For training at the Whitehead Institute with Dr. David Sabatini. Tuning Translation to Environmental Changes.

Alexander P. Fields, Ph.D. For training at the University of California, San Francisco with Dr. Jonathan Weissman. Systematic search for protein moonlighting functions.

Andrew J.P. Fink, Ph.D. For training at Columbia University with Dr. . Innate and learned odor representations in olfactory cortical networks.

Hidehiko Inagaki, Ph.D. For training at the HHMI Janelia Farm Research Campus with Dr. Karel Svoboda. Neuronal mechanism of choice selective short term memory in the mouse premotor cortex.

Joseph G. Jardine, Ph.D. For training at the Scripps Research Institute with Dr. Dennis Burton. Guiding rational vaccine design by antibody repertoire analysis.

Chengcheng Jin, Ph.D. For training at the Massachusetts Institute of Technology with Dr. Tyler Jacks. Dissecting the role of innate immunosurveillance in regulating pre-emergent tumor cells.

Vignesh Kasinath, Ph.D. For training at the University of California, Berkeley with Dr. Eva Nogales. Protein-RNA mediated epigenetic regulation of gene expression.

Aleksandar D. Kostic, Ph.D. For training at the Broad Institute with Dr. Ramnik Xavier. Identifying and characterizing gut microbial pathways and products that contribute to type 1 diabetes.

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James A. Kraemer, Ph.D. For training at the Massachusetts Institute of Technology with Dr. Michael Laub. Toxin-antitoxin systems coordinate the response to proteotoxic stress in E. coli.

Evgeny Kvon, Ph.D. For training at the Lawrence Berkeley National Laboratory with Drs. Axel Visel and Len Pennacchio. Functional in vivo exploration of mammalian distant- acting enhancer-promoter interactions.

Ya-tang Li, Ph.D. For training at the California Institute of Technology with Dr. Markus Meister. Neural Circuit Dynamics in Superior Colliculus Underlying Innate Defensive Behaviors.

Yanxin Liu, Ph.D. For training at the University of California, San Francisco with Dr. David Agard. Molecular mechanism of client protein remodeling by the Hsp90 molecular chaperone.

Matthew R. Lovett-Barron, Ph.D. For training at Stanford University with Dr. . Whole-brain neuromodulatory dynamics underlying internal state.

David B. Lyons, Ph.D. For training at the University of California, Berkeley with Dr. Daniel Zilberman. The Role of Histone H1 in the Regulation of Transposable Elements.

Maxim B. Prigozhin, Ph.D. For training at Stanford University with Dr. Steven Chu. Biogenesis of bacterial outer membrane vesicles.

Arthur Prindle, Ph.D. For training at the University of California, San Diego with Dr. Gurol Suel. Drug Resistance as a Collective Phenomenon.

Michal Rabani, Ph.D. For training at Harvard University with Dr. Alexander Schier. Post- transcriptional regulation during early development.

Mingxuan Sun, Ph.D. For training at the University of California, Berkeley with Dr. Carlos Bustamante. Structural Dynamics and Intersubunit Coordination of a Homomeric Ring ATPase.

Albert Tsao, Ph.D. For training at Stanford University with Dr. Mark Schnitzer. Probing the mechanisms for insight-based learning through continuous large-scale optical recordings.

Lamia A. Wahba, Ph.D. For training at Stanford University with Dr. Andrew Fire. Probing the Function and Evolution of piRNAs.

Fengzhu Xiong, Ph.D. For training at the Harvard University with Drs. Olivier Pouquie and L. Mahadevan. The physical, cellular and molecular mechanisms of symmetrical development.

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LIST OF AWARDS FOR 2015

Of the 444 applications received in 2015, 431 (98%) were from candidates with the Ph.D. or comparable degree awarded or pending, 9 from candidates with the M.D./Ph.D. degree, and 2 were from candidates with the M.D. degree. Of these applicants, 38 qualified for personal interview, and 23 were awarded fellowships.

Tslil Ast, Ph.D. For training at the Broad Institute with Dr. Vamsi Mootha. Dissecting mitochondrial heterogeneity within the cell.

Pau Creixell, Ph.D. For training at the Massachusetts Institute of Technology with Drs. Michael B. Yaffe and Rama Ranganathan. Comprehensive Characterization of Mutant Subclones Driving Oncogenic Transformation and Resistance to Therapy at Amino-Acid Level Resolution.

Kayvon P. Daie, Ph.D. For training at HHMI Janelia Farm Research Campus with Dr. Karel Svoboda. Probing network mechanisms underlying persistent activity in a short- term memory circuit.

Maria Ellen De Obaldia, Ph.D. For training at The Rockefeller University with Dr. Leslie Vosshall. Modeling mosquito attraction to human skin microbiota in mice.

Lihui Feng, Ph.D. For training at Washington University in St. Louis with Dr. Jeffrey Gordon. Characterizing the molecular mechanisms of gut microbiota development and its relevance to childhood undernutrition.

Kevin J. Forsberg, Ph.D. For training at the Fred Hutchinson Cancer Research Center with Dr. Harmit Malik. Revealing the Hidden World of CRISPR-Cas Antagonists.

Xiaojing J. Gao, Ph.D. For training at the California Institute of Technology with Dr. Michael Elowitz. Bottom-up analysis of Eph signaling in single cells and cell populations.

Andrew R. Gehrke, Ph.D. For training at Harvard University with Dr. Mansi Srivastava. The Regulatory Landscape of Whole-Body Regeneration.

Arielle D. Glatman Zaretsky, Ph.D. For training at the National Institute of Allergy and Infectious Diseases with Dr. Yasmine Belkaid. Long term impact of infection on lymphatic function and repair.

Fatma Ezgi Hacisuleyman, Ph.D. For training at the University of California, San Francisco with Dr. Jonathan Weissman. Regulation of pioneering translation through ribosome localization.

Mark A. Herzik Jr., Ph.D. For training at the Scripps Research Institute with Dr. Gabriel Lander. Interrogating the Role of Mitochondrial Protein Import Pathways in Disease Progression.

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Joshua H. Jennings, Ph.D. For training at Stanford University with Dr. Karl Deisseroth. A rapid brain-wide search for a neural clock of aging.

Ann Kennedy, Ph.D. For training at the California Institute of Technology with Dr. David Anderson. Dissecting the neural population code underlying social behaviors.

Chi Nguyen, Ph.D. For training at the Massachusetts Institute of Technology with Dr. Cathy Drennan. Flipping the Anaerobic Switch: The Structural Characterization of the E. coli Fumarate-Nitrate-Reductase Regulator (FNR) and FNR Regulatory Complexes.

Tomas Pluskal, Ph.D. For training at the Whitehead Institute with Dr. Jing-Ke Weng. A multi-omics platform for exploring and exploiting plant chemodiversity.

Teniel S. Ramikie, Ph.D. For training at McLean Hospital/Harvard Medical School with Drs. Kerry Ressler and Vadim Bolshakov. Estrogenic Regulation of PAC1Rs and its Psychoprotective Effects on Fear Behavior.

Arpiar B. Saunders, Ph.D. For training at Harvard Medical School with Dr. Steve McCarroll. Deciphering adolescent brain development using high-throughput single-cell sequencing.

Jian Shu, Ph.D. For training at the Broad Institute with Dr. . Pluripotency: A precarious balance of different lineage specifying forces.

Grzegorz Sienski, Ph.D. For training at the Whitehead Institute with Dr. Susan Lindquist. Elucidating the Role of APOE4 in Dysfunctions Underlying Alzheimer’s Disease.

Lu Sun, Ph.D. For training at Stanford University with Dr. Ben Barres. Molecular and Genetic Decoding of the Organization Principle for Central Nervous System Glia.

Xuebing Wu, Ph.D. For training at Whitehead Institute with Dr. David Bartel. Genome- wide discovery of mRNAs with noncoding functions.

Xiaofei Yu, Ph.D. For training at The Rockefeller University with Dr. Jeffrey Friedman. Remotely controlled immunotherapy for obesity-associated insulin resistance.

Olena Zhulyn, Ph.D. For training at Stanford University with Dr. Maria Barna. Ribosomes in regeneration: rapid translational activation in wound healing and repair.

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SCIENTIFIC PUBLICATIONS

The following publications based on research aided by Foundation support have appeared in the medical literature by those who completed their fellowships through December 31, 2015. Included in this listing may be articles received too late to be included in the previous report. An asterisk identifies the author supported by the Foundation.

Tomer R, Lovett-Barron M, Kauvar I, *Andalman A, Burns VM, Sankaran S, Grosenick L, Broxton M, Yang S, Deisseroth K. SPED light sheet microscopy: fast mapping of biological system structure and function. Cell. Cell 163, 1796–1806, December 17, 2015.

Yang SJ, Allen WE, Kauvar I, *Andalman AS, Young NP, Kim CK, Marshel JH, Wetzstein G, Deisseroth K. Extended field-of-view and increased-signal 3D holographic illumination with time-division multiplexing. Optics Express. 14 Dec 2015 | Vol. 23, No. 25 | DOI:10.1364/OE.23.032573.

Chung K., Wallace J., Kim S.Y., Kalyanasundaram, S., *Andalman A.S., Davidson T.J., Mirzabekov J.J., Zalocusky K.A., Mattis J., Denisin A.K., Pak S., Bernstein H., Ramakrishnan C., Grosenick L., Gradinaru V., Deisseroth K. Structural and molecular interrogation of intact biological systems. Nature. Advance Online Publication 2013 Apr 10. doi:10.1038/nature12107.

Tye KM, Mirzabekov JJ, Warden MR, Ferenczi EA, Tsai HC, Finkelstein J, Kim SY, Adhikari A, Thompson KR, *Andalman AS, Gunaydin LA, Witten IB, Deisseroth K. Dopamine neurons modulate neural encoding and expression of depression-related behaviour. Nature. Advance Online Publication 2012 Dec. doi:10.1038/nature11740.

*D Aronov, DW Tank. Engagement of Neural Circuits Underlying 2D Spatial Navigation in a Rodent Virtual Reality System. Neuron 84 (2), 442-456. 2014 Oct.

*Azim, E., Jiang, J., Alstermark, B., Jessell, T.M. (2014). Skilled reaching relies on a V2a propriospinal internal copy circuit. Nature. 508(7496): 357-363. PMCID: PMC4230338.

Fink, A.J.P., Croce, K.R., Huang, Z.J., Abbott, L.F., Jessell, T.M., *Azim, E. (2014). Presynaptic inhibition of spinal sensory feedback ensures smooth movement. Nature. 509(7498): 43-48. Cover article. PMCID: PMC4292914.

*Azim, E. (2014). Shortcuts and checkpoints on the road to skilled movement. Eppendorf and Science Prize for Neurobiology- Grand Prize essay. Science. 346(6209): 554-5. PMID: 25359954.

*Azim, E., Fink, A.J.P., Jessell, T.M. (2014). Internal and external feedback circuits for forelimb movement. Review. Cold Spring Harbor Symposia on Quantitative Biology, Volume LXXIX. 79: 81-92. PMCID: PMC4475648.

Miri, A., *Azim, E., Jessell, T.M. (2013). Edging toward entelechy in motor control. Perspective article. Neuron. 80(3): 827-834. PMID: 24183031.

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MY Pecot, W Tadros, A Nern, *M Bader, Y Chen, SL Zipursky. Multiple interactions control synaptic layer specificity in the Drosophila visual system. Neuron 77 (2), 299-310.

Ya Guo, Quan Xu, *Daniele Canzio, Jia Shou, Jinhuan Li, David U. Gorkin, Inkyung Jung, Haiyang Wu, Yanan Zhai, Yuanxiao Tang, Yichao Lu, Yonghu Wu, Zhilian Jia, Wei Li, Michael Q. Zhang, Bing Ren, Adrian R. Krainer, Tom Maniatis, and Qiang Wu. CRISPR Inversion of CTCF Sites Alters Genome Topology and Enhancer/Promoter Function. 2015, Cell 162, 900– 910

*Bicknell A, Cenik C, and Moore MJ. (2012) Introns in UTRs: Why we should stop ignoring them. BioEssays, 34(12); 1025-34.

*Brohawn, SG, Campbell, EB & MacKinnon, R. Physical mechanism for gating and mechanosensitivity of the human TRAAK K+ channel. Nature 516, 126-30 (2014).

*Brohawn, SG, Su, Z & MacKinnon, R. Mechanosensitivity is mediated directly by the lipid membrane in TRAAK and TREK1 K+ channels. Proceedings of the National Academy of Sciences 111, 3614–3619 (2014).

*Brohawn, SG, Campbell, EB & MacKinnon, R. Domain-swapped chain connectivity and gated membrane access in a Fab-mediated crystal of the human TRAAK K+ channel. Proceedings of the National Academy of Sciences 110, 2129–2134 (2013).

Flynn RA, Do BT, Rubin AJ, *Calo E, Lee B, Kuchelmeister H, Rale M, Chu C, Kool ET, Wysocka J, Khavari PA, Chang HY. 7SK-BAF axis controls pervasive transcription at enhancers. Nat Struct Mol Biol. 2016 Mar;23(3):231-8.

*Calo, E., Flynn, R. A., Martin, L., Spitale, R. C., Chang, H. Y., Wysocka, J. RNA helicase DDX21 coordinates transcription and ribosomal RNA processing. Nature 2015; 518 (7538): 249- 253.

Buecker, C., Srinivasan, R., Wu, Z., *Calo, E., Acampora, D., Faial, T., Simeone, A., Tan, M., Swigut, T., Wysocka, J. Reorganization of Enhancer Patterns in Transition from Naive to Primed Pluripotency Cell Stem Cell. 2014; 14 (6): 838-853.

*Calo, E., Wysocka, J. Modification of Enhancer Chromatin: What, How, and Why? Molecular Cell. 2013; 49 (5): 825-837.

*Chang, A.J., Ortega, F.E., Riegler, J., Madison, D.V. and Krasnow, M.A. (2015) Oxygen regulation of breathing through an olfactory receptor activated by lactate. Nature 527, 240-244.

*Chang, A., Ortega, F., Riegler, J., Madison, D., and Krasnow, M.A.(2015) Oxygen control of breathing by an olfactory receptor activated by lactate. Nature, 527, 240-244.

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*Daniel J. Dickinson, Ariel M. Pani, Jennifer K. Heppert, Christopher D. Higgins, Bob Goldstein. Streamlined Genome Engineering with a Self-Excising Drug Selection Cassette. Genetics. June 3, 2015; DOI: 10.1534/genetics.115.178335.

*Doulatov S, and Daley GQ. A stem cell perspective on cellular engineering. Science 2013; 342.

*Doulatov S, Vo LT, Chou SS, Kim PG, Arora N, Li H, Bernstein ID, Collins JJ, Zon LI, and Daley GQ. Induction of multipotential hematopoietic progenitors from human pluripotent cells via respecification of myeloid precursors. Cell Stem Cell. 2013; 13(4): 459-70

*Feinberg, E.H. and Meister, M. Orientation columns in the mouse superior colliculus. (2015) Nature. 519 (7542): 229-32.

*Christopher B Ford, Jason M Funt, Darren Abbey, Luca Issi, Candace Guiducci, Diego A Martinez, Toni Delorey, Bi yu Li, Theodore C White, Christina Cuomo, Reeta P Rao, Judith Berman, Dawn A Thompson, Aviv Regev. The evolution of drug resistance in clinical isolates of Candida albicans. eLife 2015;4:e00662.

Chen BC, Legant WR, Wang K, Shao L, Milkie DE, Davidson MW, Janetopoulos C, Wu XS, Hammer JA, Liu Z, English BP, Mimori-Kiyosue Y, Romero DP, Ritter AT, Lippincott- Schwartz J, *Fritz-Laylin L, Mullins RD, Mitchell DM, Bembenek JN, Reymann AC, Böhme R, Grill SW, Wang JT, Seydoux G, Tulu US, Kiehart DP, Betzig E. Lattice light-sheet microscopy: imaging molecules to embryos at high spatiotemporal resolution. Science. 2014 Oct 24; 346(6208):1257998. PMID: 25342811; PMCID: PMC4336192.

*Hammoud SS, Low DH, Yi C, Lee CL, Oatley JM, Payne CJ, Carrell DT, Guccione E, Cairns BR. Transcription and imprinting dynamics in developing postnatal male germline stem cells. Genes Dev. 2015 Nov 1;29(21):2312-24.

*Hammoud SS, Low D, Yi C, Carrell DT, Guccione E and Cairns BR (2014). Chromatin Strategies for Maintaining Pluripotency in Adult Germline Stem Cells and Mammalian Spermatogenesis. Cell Stem Cell. 2014 Aug 7;15(2):239-53.

*Saher Sue Hammoud, Bradley R Cairns, David A Jones . Epigenetic regulation of colon cancer and intestinal stem cells. Cell Biology. Volume 25, Issue 2, April 2013, Pages 177–183.

*Richard K. Hite, Xiao Tao & Roderick MacKinnon. Structural basis for gating the high- conductance Ca2+-activated K+ channel. Nature 541, 52–57 (2017).

*Hite RK, Yuan P, Li Z, Hsuing Y, Walz T, MacKinnon R. “Cryo-EM structure of Slo2.2 Na+- activated K+ channel.” Nature. 2015. 527:198-203.

*Hite RK, Butterwick JA, MacKinnon R. Phosphatidic acid modulation of Kv channel voltage sensor function.” eLife. 2014. 3:e04366.

*K.E. Kasza, D.L. Farrell, J.A. Zallen. Spatiotemporal control of epithelial remodeling by regulated myosin phosphorylation. Proceedings of the National Academy of Sciences. 2014; 111(32):11732.

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*Lamason RL, Bastounis E, Kafai NM, Serrano R, Del Álamo JC, Theriot JA, Welch MD.Rickettsia Sca4 reduces vinculin-mediated intercellular tension to promote spread. Cell 167: 670-683.e10 (2016)

Reed SC, *Lamason RL, Risca VI, Abernathy E, Welch MD. Rickettsia actin-based motility occurs in distinct phases mediated by different actin nucleators. Curr Biol 24: 98-103 (2014)

Welch MD, Reed SC, *Lamason RL, Serio AW. Expression of an epitope-tagged virulence protein in Rickettsia parkeri using transposon insertion. PLoS One 7: e37310 (2012)

Jalal K Baruni, *Brian Lau & C Daniel Salzman. Reward expectation differentially modulates attentional behavior and activity in visual area V4. Nature Neuroscience 18, 1656–1663 (2015).

Dubey, R., *Lebensohn, A., Bahrami-Nejad, Z., Marceau, C., Champion, M., Gevaert, O., Sikic, B. I., Carette, J. E., Rohatgi, R. Chromatin-remodeling complex SWI/SNF controls multidrug resistance by transcriptionally regulating the drug efflux pump ABCB1. Cancer Res. 2016 Oct 1;76(19):5810-5821. Epub 2016 Aug 8.

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*McLaughlin, R. N., Young, J. M., Yang, L., Neme, R., Wichman, H. A. & Malik, H. S. (2014) Positive Selection and Multiple Losses of the LINE-1-derived L1TD1 Gene in Mammals Suggest a Dual Role in Genome Defense and Pluripotency. PLOS Genetics 10: e1004531.

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FINANCIAL STATEMENTS THE HELEN HAY WHITNEY FOUNDATION Grants and Fellowships Paid

1947 $56,130 1981-1982 $879,009 1948 $90,329 1982-1983 $916,871 1949 $161,252 1983-1984 $939,950 1950 $225,747 1984-1985 $971,522 1951 $248,357 1985-1986 $1,008,528 1952 $250,592 1986-1987 $1,013,578 1953 $235,463 1987-1988 $1,071,616 1954 $215,550 1988-1989 $1,189,620 1955 $207,385 1989-1990 $1,304,464 1956 $189,322 1990-1991 $1,257,586 1957 (to June 30) $74,291 1991-1992 $1,347,051 1957-1958 $190,618 1992-1993 $1,179,053 1958-1959 $327,051 1993-1994 $1,437,364 1959-1960 $302,799 1994-1995 $1,478,573 1960-1961 $313,058 1995-1996 $1,430,955 1961-1962 $345,616 1996-1997 $1,514,142 1962-1963 $366,869 1997-1998 $1,632,663 1963-1964 $411,466 1998-1999 $1,840,414 1964-1965 $450,260 1999-2000 $1,929,966 1965-1966 $455,036 2000-2001 $1,755,148 1966-1967 $490,070 2001-2002 $2,688,021 1967-1968 $498,850 2002-2003 $2,824,955 1968-1969 $542,033 2003-2004 $2,750,190 1969-1970 $526,698 2004-2005 $2,486,105 1970-1971 $700,202 2005-2006 $2,190,768 1971-1972 $618,935 2006-2007 $2,364,533 1972-1973 $579,649 2007 (to Dec 31) $777,292 1973-1974 $566,145 2008 $2,488,613 1974-1975 $552,629 2009 $2,474,765 1975-1976 $578,233 2010 $2,610,683 1976-1977 $579,417 2011 $2,949,942 1977-1978 $663,775 2012 $2,880,533 1978-1979 $606,831 2013 $2,874,901 1979-1980 $810,571 2014 $2,944,182 1980-1981 $778,602 2015 $3,457,735

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The Helen Hay Whitney Foundation Statement of Financial Position December 31, 2014

ASSETS Current Assets Cash and cash equivalents $ 4,024,087 Interest and dividends receivable 134,597 Due from broker 86,801 Prepaid excise taxes 25,000 Prepaid expenses 2,560 Total current assets 4,273,045

Investments at fair value 60,165,786 Furniture and equipment, net of accumulated depreciation of $24,044 2,946 Deposits 1,457 Total assets $ 64,443,234

LIABILITIES AND UNRESTRICTED NET ASSETS Liabilities Current Liabilities Grants and fellowships payable $ 859,625 Due to broker 30,548 Other payables 40,174 Excise taxes and filing fees payable - Total current liabilities 930,347

Deferred excise taxes payable 204,435 Total liabilities 1,134,782

Unrestricted net assets 63,308,452 Total liabilities and unrestricted net assets $ 64,443,234

COMPLETE AUDITED FINANCIAL STATEMENTS OF THE HELEN HAY WHITNEY FOUNDATION ARE AVAILABLE FOR INSPECTION AT THE FOUNDATION'S OFFICE AT 20 SQUADRON BLVD, SUITE 630, NEW CITY, NY 10956

Page 23 The Helen Hay Whitney Foundation Statement of Activities Year Ended December 31, 2014

Revenue Contributions $ 1,834,970 Interest 277,470 Dividends 649,980 Partnership income 783,003 Total revenue 3,545,423

Expenses Fellowships and grants 2,944,182 Program expenses 305,470 Fund raising expenses 5,641 General and administrative 429,351 Provision for taxes and fees 110,507 Total expenses 3,795,151

Decrease in net assets before net realized and net unrealized gains on investments (249,728)

Realized and unrealized gains on investments Net realized gains on investments 2,642,695 Net unrealized gains on investments 196,396 2,839,091

Increase in net assets 2,589,363 Unrestricted net assets, beginning of year 60,719,089 Unrestricted net assets, end of year $ 63,308,452

COMPLETE AUDITED FINANCIAL STATEMENTS OF THE HELEN HAY WHITNEY FOUNDATION ARE AVAILABLE FOR INSPECTION AT THE FOUNDATION'S OFFICE AT 20 SQUADRON BLVD, SUITE 630, NEW CITY, NY 10956

Page 24 The Helen Hay Whitney Foundation Statement of Financial Position December 31, 2015

ASSETS Current Assets Cash and cash equivalents $ 6,899,631 Interest and dividends receivable 128,793 Receivable from limited partnership investment 3,570,255 Due from broker 88,645 Prepaid excise taxes 39,000 Prepaid expenses 2,560 Total current assets 10,728,884

Investments at fair value 52,263,872 Furniture and equipment, net of accumulated depreciation of $24,724 13,664 Deposits 1,457 Total assets $ 63,007,877

LIABILITIES AND UNRESTRICTED NET ASSETS Liabilities Current Liabilities Grants and fellowships payable $ 993,624 Due to broker 140,163 Other payables 40,023 Excise taxes and filing fees payable - Total current liabilities 1,173,810

Deferred excise taxes payable 169,735 Total liabilities 1,343,545

Unrestricted net assets 61,664,332 Total liabilities and unrestricted net assets $ 63,007,877

COMPLETE AUDITED FINANCIAL STATEMENTS OF THE HELEN HAY WHITNEY FOUNDATION ARE AVAILABLE FOR INSPECTION AT THE FOUNDATION'S OFFICE AT 20 SQUADRON BLVD, SUITE 630, NEW CITY, NY 10956

Page 25 The Helen Hay Whitney Foundation Statement of Activities Year Ended December 31, 2015

Revenue Contributions $ 2,347,305 Interest 230,884 Dividends 722,298 Partnership loss (78,171) Total revenue 3,222,316

Expenses Fellowships and grants 3,457,735 Program expenses 316,724 Fund raising expenses 6,143 General and administrative 441,407 Provision for taxes and fees 59,057 Total expenses 4,281,066

Decrease in net assets before net realized and net unrealized gains (losses) on investments (1,058,750)

Realized and unrealized gains (losses) on investments Net realized gains on investments 1,728,394 Net unrealized (losses) on investments (2,313,764) (585,370)

Decrease in net assets (1,644,120) Unrestricted net assets, beginning of year 63,308,452 Unrestricted net assets, end of year $ 61,664,332

COMPLETE AUDITED FINANCIAL STATEMENTS OF THE HELEN HAY WHITNEY FOUNDATION ARE AVAILABLE FOR INSPECTION AT THE FOUNDATION'S OFFICE AT 20 SQUADRON BLVD, SUITE 630, NEW CITY, NY 10956

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TO ALL FORMER HELEN HAY WHITNEY FOUNDATION FELLOWS:

The Foundation maintains a former fellows' directory on our website at www.hhwf.org.

Please help us keep our former fellows' directory up-to-date.

Search your information at: http://hhwf.org/directory/former-fellow-search/

You may update your information at: http://hhwf.org/directory/submit-directory-listing/

or via email: [email protected]

Thank you

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