Gagna A. & Ch. Van Heck

GAGNA A. & CH. VAN HECK PRIZE 2015

18 NOVEMBER 2015

Contact:

Mr Bruno MORAUX

02/504.92.40

GAGNA A. & CH. VAN HECK PRIZE

The triennial and international « Gagna A. & Ch. Van Heck Prize », amounting to 75.000 €, rewards a scientist or a Medical Doctor, in recognition of a research work which has contributed to the treatment of a disease currently incurable, or which has raised hopes for curing the disease.

This Prize was first granted in 2003 and constitutes one of the most prestigious Belgian Prize in biomedical science.

The Prize is awarded to:

Stephen P. JACKSON

Ph.D. in Molecular Biology, University of Edinburgh (UK) Head of Cancer Research UK Laboratories, The Gurdon Institute, University of Cambridge (UK) Professor of Biology, University of Cambridge (UK)

Identification of key mechanisms for the detection, signaling and repair of DNA damages.

Professor Stephen Jackson’s pioneering academic research has provided us with many of the key principles by which cells detect DNA damage, signal its presence and mediate its repair. His discoveries have had particularly major impacts on our understanding of how human cells deal with the most toxic of all DNA damages: the DNA double-strand break (DSB). In addition to establishing some of the most important foundations of present-day DNA repair research, Jackson’s work has also shown us how cellular dysfunction and disease arise when DNA repair or DNA-damage signaling goes awry. Additionally, his work has led to the development of an innovative new approach to cancer therapy that kills cancer cells whose ability to repair DNA damage has been compromised.

MEMBERS OF THE JURY GAGNA A. & CH. VAN HECK PRIZE 2015

Mr COLIGE Alain Senior Research Associate of the F.R.S. - FNRS at Université de Liège

Mr LEO Oberdan Professor at Université Libre de Bruxelles President of the Jury

Mr LETESSON Jean-Jacques Professor at Université de Namur

Mr OCTAVE Jean-Noël Professor at Université Catholique de Louvain

Mr PARMENTIER Marc Professor at Université Libre de Bruxelles

Mrs VAN LINT Carine Research Director of the F.R.S. - FNRS at Université Libre de Bruxelles

Mr VIKKULA Miikka Professor at Université Catholique de Louvain

Stephen P. JACKSON

Cancer Research UK Laboratories The Gurdon Institute University of Cambridge, UK

“Identification of key mechanisms for the detection, signaling and repair of DNA damages”

Summary

Background

Professor Stephen Jackson’s transformational academic achievements relate to cellular events that detect, signal the presence of and repair DNA damage; events whose defects contribute to many age-related human diseases. Prior to Jackson becoming an independent researcher, it had been established that DNA damage occurs in cells. It was also known that there are certain ways of repairing different forms of DNA damage. However, at that time it was not known that DNA damage in fact triggers a much wider array of intracellular events. Also, nothing was known about the molecular nature of the major DNA repair system that responds to the most toxic of all forms of DNA damage: double-strand breaks (DSBs).

The beginning: a DNA-damage activated protein kinase (DNA-PK)

Jackson entered the DNA repair field through studying the DNA-dependent protein kinase (DNA-PK). After defining the DNA-PK target consensus (subsequently found to be similar to those of the related kinases ATM and ATR), he showed DNA-PK to comprise a catalytic subunit (DNA-PKcs) and a DNA-targeting protein (Ku). Jackson’s subsequent findings relating to DNA repair all emanated from his ensuing, seminal discovery in 1993 that DNA-PK enzyme activity is triggered by DSBs through the ability of Ku to bind to broken DNA ends. This provided a key conceptual basis for cellular DNA-damage detection, repair and signaling, and paved the way for Jackson’s many subsequent transformational impacts on our understanding of such medically important processes.

DNA-PK forms the basis for the NHEJ system of DSB repair

Soon after showing that DSBs activate DNA-PK, Jackson and colleagues established that Ku and DNA-PKcs are crucial for the pathway called non-homologous end-joining (NHEJ) that repairs most DSBs in all complex organisms, including humans. In addition to showing how Ku or DNA-PKcs loss causes cells to become hypersensitive to DNA damage triggered by agents such as ionizing radiation, Jackson’s work also showed that NHEJ proteins are needed for immune system development through mediating immune- receptor gene rearrangements by V(D)J recombination.

Jackson and colleagues subsequently identified and/or helped determine the functions of all other core human NHEJ proteins, including the DNA ligase IV-XRCC4 complex and the XRCC4-related proteins XLF and PAXX. He also established – contrary to dogma at the time – that related NHEJ components and processes are conserved in yeast and in

certain bacteria. More recently, Jackson and colleagues showed how DSB repair by NHEJ and another pathway, homologous recombination, are affected by chromatin structure, and how they are reciprocally regulated by cell-cycle status via factors such as yeast Sae2 and its human counterpart CtIP.

Other DNA-damage activated kinases, pathways and processes

By showing how DNA damage detection can induce protein post-translational modifications (PTMs), Jackson’s work on DNA-PK provided a paradigm for how DNA- damage triggers intracellular signaling events whose amplification and diversification subsequently influence myriad aspects of cellular physiology. In this regard, another landmark in DNA repair and DNA-damage signaling was Jackson’s cloning of the cDNA for DNA-PKcs. His ensuing work established that DNA-PK is a bona fide protein kinase, despite its catalytic domain being most related to those of inositol phospholipid kinases. Furthermore, his work revealed that the DNA-PKcs kinase domain has homology with those of the proteins ATM and ATR, which Jackson and others then showed work through related DNA-damage induction mechanisms. Jackson’s many contributions to this area included purifying ATM and ATR, providing assays for their enzymatic activities, identifying targets, and identifying protein partners that function to recruit/activate ATM or ATR by mechanisms analogous to the Ku-DNA-PKcs paradigm. Jackson also established that ATM- and ATR-mediated signaling events are connected to one another and cell-cycle regulated.

During his work on ATM and ATR, Jackson showed how these and additional factors are targeted to DNA-damage sites, and interact in regulated ways by phosphorylation- dependent recognition mechanisms via specialized protein structures such as FHA domains and BRCT domains. In this regard, it is noteworthy that Jackson defined the first function for a chromatin modification in cellular DSB responses by demonstrating that phosphorylation of histone H2A(X) affects cellular sensitivity to DNA damage and promotes DNA repair. He also provided a mechanism for this function in mammalian cells by establishing that it serves as a binding site for the protein MDC1, which then mediates the recruitment of many other proteins to DNA-damage sites.

Jackson’s work also provided us with insights into how DNA repair is affected by diverse other cellular factors and, conversely how DNA repair and DNA-damage signaling impinge on various aspects of cellular biology. For example, he showed that DSB-repair proteins control normal telomere functions. He also discovered that mammalian cell replicative senescence – the arrest of cell proliferation upon repeated passage of primary human cells in culture – reflects an ATM/ATR based DNA-damage response triggered by shortened telomeres. More recently, Jackson’s research established how other PTMs, such as ubiquitylation, sumoylation, poly-ADP ribosylation, acetylation and methylation play key roles in cellular responses to DSBs.

A new therapeutic approach: killing cancer cells by “synthetic lethality”

By working in yeast, Jackson found that defects in some DNA-repair components only 6

cause marked DNA-damage hypersensitivity in the absence of other DNA repair factors, a phenomenon based on the genetic principle termed “synthetic lethality”. Recognizing that some cancer cells lack certain DNA repair factors and would thus likely be particularly reliant on other DNA-repair pathways, Jackson realized that “synthetic lethal” relationships between pathways might also be exploited pharmacologically and clinically.

Through refining biochemical assays for enzymes such as DNA-PK, ATM and ATR and the DNA-damage activated DNA-repair enzyme PARP in his academic laboratory in the mid-1990s, Jackson found that it was possible to identify small-molecule compounds that could inhibit such enzymes. Based on this and his prior yeast and mammalian cell findings, Jackson founded the company, KuDOS Pharmaceuticals, and as part-time Chief Scientific Officer (CSO), he steered KuDOS to become an integrated drug- discovery and development company. KuDOS was acquired in 2006 by AstraZeneca but functioned as an AstraZeneca subsidiary, with Jackson as CSO until 2010. Based on his work showing how other PTMs such as ubiquitylation and sumoylation control key DNA repair processes, in mid-2011 Jackson founded the company MISSION Therapeutics to develop drugs targeting enzymes of the ubiquitin PTM system to treat cancer via synthetic-lethality and other mechanisms.

The programmes Jackson established and steered within KuDOS produced compounds such as ATM, DNA-PK and PARP inhibitors that are now widely used as research tools. Moreover, the KuDOS-generated compounds olaparib (a PARP1/2, inhibitor), AZD8055 (an mTOR inhibitor) and AZ20 (an ATR inhibitor) are now in clinical trials. Most notably, Jackson and colleagues showed that olaparib and related PARP inhibitors exhibit striking cytotoxicity towards cancer cells impaired in homologous-recombination through them possessing mutations in BRCA1 or BRCA2, thus validating the “synthetic-lethal pharmacology concept” that Jackson had outlined in his original 1997 KuDOS Business Plan. This nurtured the development of olaparib as the first of a new class of anti-cancer agents working via synthetic lethality: killing cancer cells by targeting their genetic dependency on particular DNA repair systems (their “Achilles’ heels”) but having little effect on normal cells and therefore having few toxic side-effects.

Olaparib (marketed by AstraZeneca as LynparzaTM) recently received approval by the US FDA (Food and Drug Administration) and the EMA (European Medicines Agency) for treating ovarian cancers with BRCA1/2 mutations. There is now much excitement that olaparib and other PARP inhibitors may soon become approved for treating various other cancers.

Curriculum Vitae

Stephen Philip JACKSON, FRS, FMedSci

Frederick James Quick and Cancer Research UK Professor of Biology,

Head of Cancer Research UK Laboratories,

The Gurdon Institute, University of Cambridge, UK

PERSONAL DETAILS

Date of Birth: July 17, 1962 (age 52)

Place of Birth: Nottingham, UK

Nationality: British

Work Address: The Gurdon Institute University of Cambridge Tennis Court Road Cambridge, CB2 1QN, UK

Telephone: (01223) 334102 or 331725

Fax: (01223) 334089

E-mail: [email protected]

PROFESSIONAL EXPERIENCE

2012-present Associate Faculty Member, The Sanger Institute, Hinxton, Cambridge, UK

2011-present Founding Scientist and Chief Scientific Officer (part-time), MISSION Therapeutics Ltd., Babraham, Cambridge, UK.

2010-present Recipient of salary supplement from Cancer Research UK

2009-present Frederick James Quick and Cancer Research UK Professor of Biology, Department of Biochemistry, University of Cambridge, UK.

2004-present Head, Cancer Research UK Laboratories, The Wellcome Trust and Cancer Research UK Gurdon Institute, University of Cambridge, UK.

1995-present Senior Group Leader, The Gurdon Institute, University of Cambridge, UK.

1995-present Fellow of St John’s College, University of Cambridge, UK.

1995-2009 Frederick James Quick Professor of Biology, Department of Zoology, University of Cambridge, UK.

1997-2008 Chief Scientific Officer (part time), KuDOS Pharmaceuticals Ltd., Cambridge, UK.

1997 Sole Founding Scientist, KuDOS Pharmaceuticals Ltd., Cambridge, UK.

2001-2004 Deputy Director, The Gurdon Institute, University of Cambridge, UK.

1991-1995 Junior Group Leader, The Wellcome Trust and Cancer Research Campaign Institute, University of Cambridge, UK.

1987-1991 Postdoctoral research in the laboratory of Prof. R. Tjian, University of California at Berkeley, USA.

1985-1987 Graduate research in laboratory of Prof. J. D. Beggs, University of Edinburgh, UK.

1983-1985 Graduate research in the laboratory of Prof. J. D. Beggs, Department of Biochemistry, Imperial College of Science and Technology, London, UK.

EDUCATION

1983-1987 University of Edinburgh, Scotland, UK; Ph.D. in Molecular Biology

1980-1983 University of Leeds, UK; B.Sc. Honours (1st Class) in Biochemistry

1978-1980 High Pavement Sixth-Form College, Nottingham, UK.

1973-1978 Haywood Comprehensive School, Nottingham, UK.

AWARDS AND MEMBERSHIPS OF SCIENTIFIC ORGANIZATIONS

2015 The Gagna A. & Ch. Van Heck Prize “For determining how cells recognize and repair DNA damage and translating this knowledge towards new cancer therapies”

2015 The Sackler Lecture (delivered in Cambridge and at the Academy of Medical Sciences, London)

2014 Thomson Reuters Highly Cited Researcher

2014 Fellowship of the European Academy of Cancer Sciences

2014 Fellowship of Imperial College Faculty of Medicine

2012 Member of the UK BioIndustry Association

2012 Among the most highly cited European cancer researchers, 1998-2009 http://www.labtimes.org/labtimes/issues/lt2012/lt01/lt_2012_01_40_42.pdf

2011 Royal Society Buchanan Medal for distinguished contributions to the medical sciences

2011 Medical Research Council Keynote Prize Lecture; Society of Toxicology Anniversary Annual Meeting, Washington, USA. March 9th, 2011.

2010 Among the most highly cited European molecular biologists, 1997-2008 (http://www.labtimes.org/labtimes/issues/lt2010/lt05/lt_2010_05_46_48.pdf).

2009 BBSRC Innovator of the Year

2008 Elected to the Fellowship of the Royal Society

2008 Biochemical Society GlaxoSmithKline Award

2007 Honorary degree of Doctor of Science; University of Nottingham

2006 Degree of Doctor of Science; University of Cambridge

2006-present ISI Highly Cited Researchers in ISIHighlyCited.com

2005-present Member of the British Association for Cancer Research

2005-present Member of the European Association for Cancer Research

2002 Anthony Dipple Carcinogenesis Young Investigator award

2001 Elected to the Academy of Medical Sciences

1997 Elected to the European Molecular Biology Organisation

1997 Biochemical Society Colworth Medal

1997 Tenovus Medal for Cancer Research

1995 Eppendorf-Nature European Young Investigator Award

1993-present Member of the Biochemical Society

1993-present Member of the Cambridge Philosophical Society

COMMITTEE MEMBERSHIPS

Sept 2015- Wellcome Trust Collaborative Awards Committee

2013-present International Scientific Advisory Board; Netherlands Cancer Institute

2013-present Scientific Advisory Board; MRC Clinical Sciences Centre

2012-present Chair, Gurdon Institute Biological Safety Committee

2012-present Scientific Advisory Board; MRC Protein Phosphorylation and Ubiquitylation Unit, Dundee

2012-present Cancer Research UK Science Committee

2012-2013 Scientific Advisory Group; Cancer Research UK London Research Institute

2011-present Scientific Advisory Board; MRC Toxicology Unit, Leicester

2011-present Steering Committee, Cambridge Cancer Centre

2010-2012 Cancer Research UK Drug Discovery Advisory Group

2009-2013 Chairman of the Board, Scottish Centre for Cell Signalling (SCILLS), Dundee, Scotland

2008-present Scientific Advisory Board; Beatson Institute, Glasgow, Scotland

2005-present Scientific Advisory Board; Radiation Oncology & Biology Institute, Oxford

2003-present Gurdon Institute Management Committee

2009-2011 Strategic Board of the Drug Discovery Program, IFOM-IEO, Milan, Italy

STATEMENT OF RESEARCH INTERESTS

The aim of my academic research is to better understand how cells detect DNA damage and signal its presence to the DNA repair and cell cycle machineries. Towards this end, my laboratory is using a broad range of techniques and approaches, in both mammalian and yeast cells. It is my belief that a deeper knowledge of these pathways will yield a better understanding of the diseases that can arise when such pathways are lost – such as hereditary and sporadic cancer, neurodegeneration, developmental defects, immune deficiencies, infertility and premature ageing – and will suggest new strategies for treating such diseases. Much research in my lab has arisen from the paradigm of DNA damage detection and signalling that I established for the DNA-PK protein kinase a number of years ago. Although a considerable amount of my lab’s current work is still focused on pathways controlled by DNA-PK and the related kinases ATM and ATR, our research is increasingly determining how other post-translational modifications, such as protein acetylation, poly(ADP)-ribosylation, ubiquitylation and sumoylation, also control key DDR events. Further details about my academic laboratory can be found at: http://www2.gurdon.cam.ac.uk/~jacksonlab/

KUDOS PHARMACEUTICALS LTD. AND MISSION THERAPEUTICS LTD

I founded KuDOS in 1997 to translate knowledge of DNA repair and DNA-damage response (DDR) pathways into new treatments for cancer. The overall concept was to

develop drugs for use as stand-alone agents (acting via synthetic-lethality and related mechanisms) and/or to enhance the efficacy of DNA-damaging anti-cancer treatments such as chemotherapies and radiotherapy. Acting as Chief Scientific Officer (CSO) I with help from my many colleagues, took the company through three rounds of venture- capital financing, raising ~£43 million. This allowed KuDOS to develop into a fully integrated drug-discovery and drug-development company. Having generated several drug candidates and taken these to various stages of development, KuDOS was acquired in 2005/6 by AstraZeneca for $210 million; and until 2010, KuDOS functioned as a Cambridge-based subsidiary of AstraZeneca with me as CSO. The programmes that I established and steered within KuDOS produced compounds such as ATM, DNA-PK and PARP inhibitors that are now widely used as research tools. Moreover, the KuDOS- generated compound LynparzaTM/olaparib (a PARP1/2, inhibitor) is a registered drug (see below), while AZD8055 (an mTOR inhibitor) and AZ20 (an ATR inhibitor) are in clinical trials (DNA-PK and ATM inhibitors are in preclinical development).

In 2002, I initiated a collaboration between KuDOS and the laboratory of Professor Alan Ashworth (Institute for Cancer Research, London, UK) to explore the impact of DNA repair inhibitors on cells mutated in the breast and ovarian cancer predisposition genes, BRCA1 and BRCA2. This and ensuing preclinical and clinical findings revealed olaparib and related PARP inhibitors to exhibit striking cytotoxicity towards cells with BRCA1 or BRCA2 deficiencies, thus validating the “synthetic-lethal pharmacology concept” that I had in my 1997 KuDOS Business Plan: killing cancer cells by targeting their genetic dependency on particular DNA repair systems (their “Achilles’ heels”) but having little effect on normal cells and therefore having few toxic side-effects. In addition to extending lives (in some cases for many years) of cancer-trial patients, clinical data have highlighted olaparib’s potential in BRCA1/2 hereditary and non-hereditary cancers, including breast, pancreatic, prostate and ovarian. LynparzaTM/olaparib recently received FDA and EMA approval for use in ovarian cancers with BRCA1/2 mutations. LynparzaTM/olaparib is thus the first-in-class drug targeting PARP, and is the first drug to exploit the synthetic-lethality concept. It is currently being evaluated in seven Phase III trials.

Realizing that therapeutic opportunities arising from my research were not being adequately exploited, I conceived MISSION Therapeutics. MISSION has received strong venture-capital backing and set up its operations on the Babraham Science Campus near Cambridge, with me serving as its part-time Chief Scientific Officer (http://www.missiontherapeutics.com/).

SEMINARS

Over 310 invited seminars since 1992.

EDITORIAL BOARDS

2015-present Oncotarget

2012-present Science Signaling (Board of Reviewing Editors)

2010-present Biomolecules

2008-present Aging

2003-present Genes and Development

2001-present DNA Repair

1999-present EMBO Journal

1999-present Carcinogenesis

2005-present The Scientist

2004-2014 Current Biology

2011-2014 PLoS Biology

2000-2011 Nature Reviews

2001-2010 Science

2000-2010 EMBO Reports

2001-2009 Faculty of 1000

2000-2005 British Journal of Cancer

PATENTS

Granted patent applications in various territories relating to: DNA damage response (DDR) proteins as targets for anti-cancer therapies; assays for identifying small-molecule inhibitors of DDR proteins; the development and use of specific DDR inhibitors; and the demonstration of synthetic-lethality as an anti-cancer strategy based on the use of DDR inhibitors such as PARP inhibitors to selectively kill cancer cells deficient in homologous recombination mediated DNA repair (US2005227919).

MEETINGS AND CONFERENCES ORGANIZED OR CO-ORGANIZED

2016 Abcam conference “Maintenance of Genome Stability”, Panama

2015 EMBO conference “The DNA Damage Response in Cell Physiology and Disease”, Greece

2014 Abcam conference “Maintenance of Genome Stability”, St. Kitts

2013 EMBO conference “The DNA Damage Response in Cell Physiology and Disease”, Greece

2013 Keystone Symposium “Genomic Instability and DNA Repair”, Banff Canada

2012 Abcam conference “Maintenance of Genome Stability”, The Bahamas

2011 Chromosome Structure, Damage and Repair, Anavyssos, Greece

2010 A-T Workshop “Molecular Basis of A-T”, California, USA

2010 “Telomeres and Genome Stability” Conference, Marseilles, France

2010 Abcam conference “Maintenance of Genome Stability”, Antigua

2009 DNA Repair Integrated Project Symposium, Crete

2008 Abcam conference “Maintenance of Genome Stability”, Puerto Vallarta, Mexico.

2008 “Telomeres and Genome Stability” Conference, Villars-sur-Ollon, Switzerland

2006 “Telomeres and Genome Stability” Conference, Villars-sur-Ollon, Switzerland

2004 EMBO/58th Harden Conference “Telomeres and Genome Stability” Cambridge, UK. This has evolved to a regular meeting, occurring every two years

2002 International symposium “Signalling the Future” Liverpool, UK

1997 Biochemical Society Annual Symposium “Cellular Responses to Stress” Dundee, UK

1996-2005 Annual Molecular Biology and Cancer Network Conference “Genes and Cancer”, University of Warwick, UK

1995 UK-RSA Symposium “Cell Growth Control” Cape Town, South Africa

1994 The first “EMBL Conference on Gene Transcription”, Heidelberg, Germany

1993 Biochemical Society Meeting “Signalling from the plasma membrane to the Nucleus” Sheffield, UK

TEACHING (University of Cambridge)

Lecturer and examiner (2002-2013): Part IB ‘Cell and Developmental Biology’

Organizer, lecturer and examiner: Part II course ‘Cell growth and genome stability’

Senior Examiner (2004-2007): Part IB ‘Cell and Developmental Biology’

RESEARCH TRAINEES

Predoctoral: 19 Pre-doctoral students including:

1993-1994 Peter Baumann Investigator, Stowers Institute for Medical Research, Kansas, USA.

1994-1997 David Gell Menzies Research Institute, Australia

1994-1998 Simon Boulton Group Leader, Cancer Research UK, London Research Institute, London, UK.

1997-2001 Andrew McAinsh Centre for Mechanochemical Cell Biology, Warwick Medical School, University of Warwick

1997-2001 Damien D’Amours Group Leader, Institute for Research in Immunology and Cancer, University of Montreal, Canada

1999-2003 Rajat Roy Research Associate, Imperial College

1999-2003 Ali Jazayeri Head of Protein Engineering, Heptares Therapeutics, Hertfordshire

2000-2004 Philip Reaper Research Scientist with Vertex Pharmaceuticals Inc., Oxfordshire

2002-2006 Peter Ahnesorg Medical Manager, Oncology Department, Roche Switzerland

2005-2008 Robert Driscoll Postdoctoral Fellow, K. Cimprich Laboratory, Stanford, California, USA.

2006-2010 J. Ross Chapman Group Leader, University of Oxford

2006-2011 Jorrit Tjeertes Postdoctoral Fellow, Novartis, Switzerland

2010-2014 Jessica Brown SpR in Medical Oncology, Addenbrooke’s Hospital, Cambridge.

Postdoctoral: 22 Post-doctoral fellows including:

1991-1995 Robert White Professor of Biochemistry, University of York

1994-1997 Susan Critchlow Senior Staff Scientist Astra Zeneca, UK.

1994-1999 Graeme Smith Director of Research, Astra Zeneca, UK

1995-1998 Ugur Yavuzer Associate Professor, Istanbul Technical University, Istanbul, Turkey.

1996-1997 Tonya Bliss Research Scientist, Stanford University, California, USA.

1996-2002 Jessica Downs Reader in Genome Stability, MRC Genome Damage and Stability Centre, Sussex University

1996-1999 Raimundo Freire Staff Scientist, Hospital Universitario de Canarias, Tenerife, Spain

1997-2001 Steve Bell Professor of Microbiology, Sir William Dunn School of Pathology, Oxford

1997-2001 Daniel Durocher Group Leader, Samuel Lunenfeld Research

Institute Toronto, Canada

1997-2002 John Rouse Professor of Chromosome Biology and Post- Graduate Programme Coordinator, MRC Protein Phosphorylation Unit, University of Dundee, UK.

1997-2003 Fabrizio d’Adda di Fagagna Group leader, FIRC Institute for Molecular Oncology in Milan, Italy

1998-2001 Nick Lakin Lecturer, Biochemistry Department, University of Oxford, UK.

1999-2001 Brandi Williams Center for Translation Medicine at the Moran Eye Center, University of Utah, USA.

1998-2003 Soo-Hwang Teo Chief Executive, Cancer Research Initiatives Foundation, University Malaya Medical Centre, Malaysia

1999-2004 Michal Goldberg Lecturer, The Hebrew University, Jerusalem, Israel

2000-2004 Muriel Grenon Postdoctoral fellow, National University of Ireland, Galway, Ireland

2001-2004 Veronique Smits Hospital Universitario de Canarias, Tenerife, Spain

2001-2005 Manuel Stucki Group Leader, University of Zurich, Switzerland

2003-2005 Jacob Falck Novo Nordisk Biotech Fund, Denmark

2001-2006 Abdel Moumen Head Biotechnology R&D Projects, MAScIR Medical Biotechnology, Morocco

2003-2007 Alessandro Sartori Institute of Molecular Cancer Research, University of Zurich

2002-2008 Serge Gravel Sherbrooke University, Quebec, Canada

2004-2010 Pablo Huertas Group Leader in the Andalusian Center for Molecular Biology and Regenerative Medicine, University of Seville

2007-2010 Sonja Flott Business Development Associate, Abcam, Cambridge, UK.

2006-2010 Sophie Polo Group Leader, Institut Curie, Paris, France.

2006-2011 Kyle Miller Assistant Professor, University of Texas at Austin, USA

2005-2011 Jeanine Harrigan Senior Scientist, MISSION Therapeutics Ltd, Cambridge, UK.

2011-2014 Jon Travers MedImmune, Cambridge, UK.

Additional Mentoring (current):

Shikang Liang (Department of Biochemistry): PhD Mentoring Scheme

Felipe Cortes Ledesma (CSIC, Spain): EMBO Mentor

Andrej Alendar (Gurdon Institute): Postdoctoral Mentoring Scheme

Tanay Ghosh (Gurdon Institute): Postdoctoral Mentoring Scheme

Marta Teperek-Tkacz (Gurdon Institute): Postdoctoral Mentoring Scheme

Ka Hing Che (Gurdon Institute): Postdoctoral Mentoring Scheme

Joanna Loizou (CeMM, Austria): Advisor/Mentor

Saskia Suijkerbuijk (Gurdon Institute): Postdoctoral Mentoring Scheme

Claudia Wurzenberger (Gurdon Institute): Postdoctoral Mentoring Scheme

Nina Oberbeck (MRC LMB): PhD Second Supervisor

Sam Behjati (Sanger Institute): Wellcome Trust Clinical PhD External Advisor

Charlotte Sutherell (Department of Chemistry): PhD Mentoring Scheme

Therese Anderson (Gurdon Institute): Postdoctoral Mentoring Scheme

Mark Jackman (Gurdon Institute): Postdoctoral Mentoring Scheme

Alejandra Gariol (Gurdon Institute): Postdoctoral Mentoring Scheme

Bernhard Strauss (Gurdon Institute): Postdoctoral Mentoring Scheme

Antonio Campos Caro (Gurdon Institute): Postdoctoral Mentoring Scheme

Johanna Rees (Gurdon Institute): Postdoctoral Mentoring Scheme

Kei Miyamoto (Gurdon Institute): Postdoctoral Mentoring Scheme

Qiang Wu (Gurdon Institute): Postdoctoral Mentoring Scheme

Mikkel Christensen (Gurdon Institute): Postdoctoral Mentoring Scheme

PUBLICATIONS

 180 peer-reviewed research publications plus 39 peer-reviewed review articles and 24 other review articles and book chapters.

 16 first author publications.

 159 corresponding/co-corresponding author publications.

 Publications include 13 in Cell, 11 in Science, 13 in Nature, 4 in Nature Genetics, 3 in Nature Cell Biology, 1 in Nature Chemical Biology, 1 in Nature Reviews Cancer, 13 in Genes & Development, 13 in Molecular Cell, 13 in EMBO J., 13 in PNAS, 10 in Current Biology, 5 in EMBO Reports, 2 in The Journal of Cell Biology, and 11 in J. Biological Chemistry.

 h-index: 114 (Google Scholar)

 Total citations: >51,000

 i10 index: 244

Research Articles:

1. Williams, L., McDonald, C., Jackson, S., McIntosh, E. and Higgins, S. (1983). Isolation and characterization of genomic and cDNA clones for an androgen-regulated secretory protein of rat seminal vesicles. Nucleic Acids Research 11, 5021-5036.

2. Lossky, M., Anderson, G. J., Jackson, S. P. and Beggs, J. (1987). Identification of a yeast snRNP protein and detection of snRNP-snRNP interactions. Cell 51, 1019-1026.

3. Jackson, S. P. and Tjian, R. (1988). O-glycosylation of eukaryotic transcription factors implications for mechanisms of transcriptional regulation. Cell 55, 125-133.

4. Jackson, S. P., Lossky, M. and Beggs, J. D. (1988). Cloning of the RNA8 gene of Saccharomyces cerevisiae, detection of the RNA8 protein, and demonstration that it is essential for nuclear pre-mRNA splicing. Mol Cell Biol 8, 1067-1075.

5. Courey, A. J., Holtzman, D. A., Jackson, S. P. and Tjian, R. (1989). Synergistic activation by the glutamine-rich domains of human transcription factor Sp1. Cell 59, 827-836.

6. Jackson, S. P. and Tjian, R. (1989). Purification and analysis of RNA polymerase II transcription factors by using wheat germ agglutinin affinity chromatography. Proc Natl Acad Sci USA 86, 1781-1785.

7. Jackson, S. P., Macdonald, J. J., Lees-Miller, S. and Tjian, R. (1990). GC box binding induces phosphorylation of Sp1 by a DNA-dependent protein kinase. Cell 63, 155- 165.

8. Saffer, J. D., Jackson, S. P. and Thurston, S. J. (1990). SV40 stimulates expression of the trans-acting factor Sp1 at the mRNA level. Genes and Development 4, 659-666.

9. Li, R., Knight, J. D., Jackson, S. P., Tjian, R. and Botchan, M. R. (1991). Direct interaction between Sp1 and the BPV enhancer E2 protein mediates synergistic activation of transcription. Cell 65, 493-505.

10. Mastrangelo, I. A., Courey, A. J., Wall, J. S., Jackson, S. P. and Hough, P. V. C. (1991). DNA looping and Sp1 multimer links a mechanism for transcriptional synergism and enhancement. Proc Natl Acad Sci USA 88, 5670-5674.

11. Saffer, J. D., Jackson, S. P. and Annarella, M. B. (1991). Developmental expression of Sp1 in the mouse. Mol Cell Biol 11, 2189-2199.

12. Su, W., Jackson, S., Tjian, R. and Echols, H. (1991). DNA looping between sites for transcriptional activation: self-association of DNA-bound Sp1. Genes and Development 5, 820-826.

13. White, R. J., *Jackson, S. P. and Rigby, P. W. J. (1992). A role for the TATA-box-binding protein component of the transcription factor IID complex as a general RNA polymerase III transcription factor. Proc Natl Acad Sci USA 89, 1949-1953. *corresponding author.

14. White, R. J. and Jackson, S. P. (1992). Mechanism of TATA-binding protein recruitment to a TATA-less class III promoter. Cell 71, 1041-1053.

15. White, R. J., Rigby, P. W. J. and Jackson, S. P. (1992). The TATA-binding protein is a general transcription factor for RNA polymerase III. Journal of Cell Science, 1-7.

16. Bannister, A. J., Gottlieb, T. M., Kouzarides, T. and Jackson, S. P. (1993). c-Jun is phosphorylated by the DNA-dependent protein kinase in vitro; definition of the minimal kinase recognition motif. Nucleic Acids Research 21, 1289-1295.

17. Gottlieb, T. M. and Jackson, S. P. (1993). The DNA-dependent protein kinase: requirement for DNA ends and association with Ku antigen. Cell 72, 131-142.

18. Khoo, B., Brophy, B. and Jackson, S. P. (1994). Conserved functional domains of the RNA polymerase III general transcription factor BRF. Genes and Development 8, 2879-2890.

19. Rowlands, T., Baumann, P. and Jackson, S. P. (1994). The TATA-binding protein: a general transcription factor in eukaryotes and archaebacteria. Science 264, 1326- 1329.

20. Taccioli, G. E., Gottlieb, T. M., Blunt, T., Priestley, A., Demengeot, J., Mizuta, R., Lehmann, A. R., Alt, F. W., Jackson, S. P. and Jeggo, P. A. (1994). Ku80: product of the XRCC5 gene and its role in DNA repair and V(D)J recombination. Science 265, 1442-1445.

21. White, R. J., Khoo, B. C.-E., Inostroza, J. A., Reinberg, D. and Jackson, S. P. (1994). Differential regulation of RNA polymerase I, polymerase II, and polymerase III by the TBP-binding repressor Dr1. Science 266, 448-450.

22. Blunt, T., Finnie, N. J., Taccioli, G. E., Smith, G. C. M., Demengeot, J., Gottlieb, T. M., Mizuta, R., Varghese, A. J., Alt, F. W., Jeggo, P. A. and Jackson, S. P. (1995). Defective DNA-dependent protein kinase activity is linked to V(D)J recombination and DNA repair defects associated with the murine SCID mutation. Cell 80, 813–823.

23. Blunt, T., Taccioli, G. E., Priestley, A., Hafezparast, M., McMillan, T., Liu, J., Cole, C. C., White, J., Alt, F. W., Jackson, S. P., Schurr, E., Lehmann, A. R. and Jeggo, P. A. (1995). A YAC contig encompassing the XRCC5 (Ku80) DNA repair gene and complementation of defective cells by YAC protoplast fusion. Genomics 30, 320-328.

24. Finnie, N. J., Gottlieb, T. M., Blunt, T., Jeggo, P. A. and Jackson, S. P. (1995). DNA- dependent protein kinase activity is absent in xrs-6 cells: implications for site-specific recombination and DNA double-strand break repair. Proc Natl Acad Sci USA 92, 320- 324.

25. Hartley, K. O., Gell, D., Smith, G. C. M., Zhang, H., Divecha, N., Connelly, M. A., Admon, A., Lees-Miller, S. P., Anderson, C. W. and Jackson, S. P. (1995). DNA- dependent protein kinase catalytic subunit: a relative of phosphatidylinositol 3- kinase and the ataxia telangiectasia gene product. Cell 82, 849-856.

26. Hodges, P. E., Jackson, S. P., Brown, J. D. and Beggs, J. D. (1995). Extraordinary sequence conservation of the PRP8 splicing factor. Yeast 11, 337-342.

27. Kuhn, A., Gottlieb, T. M., *Jackson, S. P. and Grummt, I. (1995). DNA-dependent protein kinase: a potent inhibitor of transcription by RNA polymerase I. Genes and Development 9, 193-203. * Sole corresponding author.

28. Miller, R. D., Hogg, J., Ozaki, J. H., Gell, D., Jackson, S. P. and Riblet, R. (1995). Gene for the catalytic subunit of mouse DNA-dependent protein kinase maps to the scid locus. Proc Natl Acad Sci USA 92, 10792-10795.

29. Qureshi, S. A., Khoo, B., Baumann, P. and Jackson, S. P. (1995). Molecular cloning of the transcription factor TFIIB homolog from Sulfolobus shibatae. Proc Natl Acad Sci USA 92, 6077-6081.

30. Qureshi, S. A., Baumann, P., Rowlands, T., Khoo, B. and Jackson, S. P. (1995). Cloning and functional analysis of the TATA binding protein from Sulfolobus shibatae. Nucleic Acids Research 23, 1775-1781.

31. Sipley, J. D., Menninger, J. C., Hartley, K. O., Ward, D. C., Jackson, S. P. and Anderson, C. W. (1995). Gene for the catalytic subunit of the human DNA-activated protein kinase maps to the site of the XRCC7 gene on chromosome 8. Proc Natl Acad Sci USA 92, 7515-7519.

32. White, R. J., Gottlieb, T. M., Downes, C. S. and Jackson, S. P. (1995). Mitotic regulation of a TATA-binding-protein-containing complex. Mol Cell Biol 15, 1983-1992.

33. White, R. J., Gottlieb, T. M., Downes, C. S. and Jackson, S. P. (1995). Cell-cycle regulation of RNA polymerase III transcription. Mol Cell Biol 15, 6653-6662.

34. Baumann, P. and Jackson, S. P. (1996). An archaebacterial homologue of the essential eubacterial cell division protein FtsZ. Proc Natl Acad Sci USA 93, 6726-6730.

35. Blunt, T., Gell, D., Fox, M., Taccioli, G. E., Lehmann, A. R., Jackson, S. P. and Jeggo, P. A. (1996). Identification of a nonsense mutation In the carboxyl-terminal region of DNA-dependent protein kinase catalytic subunit in the scid mouse. Proc Natl Acad Sci USA 93, 10285-10290.

36. Boulton, S. J. and Jackson, S. P. (1996). Identification of a Saccharomyces cerevisiae Ku80 homologue: roles in DNA double-strand break rejoining and in telomeric maintenance. Nucleic Acids Research 24, 4639-4648.

37. Boulton, S. J. and Jackson, S. P. (1996). Saccharomyces cerevisiae Ku70 potentiates illegitimate DNA double-strand break repair and serves as a barrier to error-prone DNA repair pathways. EMBO Journal 15, 5093-5103.

38. DeDecker, B. S., O'Brien, R., Fleming, P. J., Geiger, J. H., Jackson, S. P. and Sigler, P. B. (1996). The crystal structure of a hyperthermophilic archaeal TATA-box binding protein. Journal of Molecular Biology 264, 1072-1084.

39. Grawunder, U., Finnie, N., Jackson, S. P., Riwar, B. and Jessberger, R. (1996). Expression of DNA-dependent protein kinase holoenzyme upon induction of lymphocyte differentiation and V(D)J recombination. European Journal of Biochemistry 241, 931-940.

40. Jongmans, W., Artuso, M., Vuillaume, M., Bresil, H., Jackson, S. P. and Hall, J. (1996). The role of Ataxia telangiectasia and the DNA-dependent protein kinase in the p53- mediated cellular response to ionising radiation. Oncogene 13, 1133-1138.

41. Lakin, N. D., Weber, P., Stankovic, T., Rottinghaus, S. T., Taylor, A. M. R. and Jackson, S. P. (1996). Analysis of the ATM protein in wild type and ataxia telangiectasia cells. Oncogene 13, 2707-2716.

42. Le Romancer, M., Cosulich, S. C., Jackson, S. P. and Clarke, P. R. (1996). Cleavage and inactivation of DNA-dependent protein kinase catalytic subunit during apoptosis in Xenopus egg extracts. Journal of Cell Science 109, 3121-3127.

43. Nicolas, N., Finnie, N. J., Cavazzanacalvo, M., Papadopoulo, D., Ledeist, F., Fischer, A., Jackson, S. P. and Devillartay, J. P. (1996). Lack of detectable defect in DNA double-strand break repair and DNA-dependent protein kinase activity in radiosensitive human severe combined immunodeficiency fibroblasts. European Journal of Immunology 26, 1118-1122.

44. Song, Q., Lees-Miller, S. P., Kumar, S., Zhang, N., Chan, D. W., Smith, G. C. M., Jackson, S. P., Alnemri, E. S., Litwack, G., Khanna, K. K. and Lavin, M. F. (1996). DNA- dependent protein kinase catalytic subunit: a target for an ICE-like protease in apoptosis. EMBO Journal 15, 3238-3246.

45. Song, Q., Burrows, S. R., Smith, G., Lees-Miller, S. P., Kumar, S., Chan, D. W., Trapani, J. A., Alnemri, E., Litwack, G., Lu, H., Moss, D. J., Jackson, S. and Lavin, M. F. (1996). Interleukin-1-b-converting enzyme-like protease cleaves DNA-dependent protein kinase in cytotoxic T cell killing. Journal of Experimental Medicine 184, 619-626.

46. White, R. J., Trouche, D., Martin, K., Jackson, S. P. and Kouzarides, T. (1996). Repression of RNA polymerase III transcription by the retinoblastoma protein. Nature 382, 88-90.

47. Williams, R. D., Lee, B. A., Jackson, S. P. and Proudfoot, N. J. (1996). Activation domains of transcription factors mediate replication dependent transcription from a minimal HIV-1 promoter. Nucleic Acids Research 24, 549-557.

48. Critchlow, S. E., Bowater, R. P. and Jackson, S. P. (1997). Mammalian DNA double- strand break repair protein XRCC4 interacts with DNA ligase IV. Current Biology 7, 588-598.

49. Larminie, C. G. C., Cairns, C. A., Mital, R., Martin, K., Kouzarides, T., Jackson, S. P. and White, R. J. (1997). Mechanistic analysis of RNA polymerase III regulation by the retinoblastoma protein. EMBO Journal 16, 2061-2071.

50. Morrison, C., Smith, G. C. M., Stingl, L., Jackson, S. P., Wagner, E. F. and Wang, Z.-Q. (1997). Genetic interaction between PARP and DNA-PK in V(D)J recombination and tumorigenesis. Nature Genetics 17, 479-482.

51. Qureshi, S. A., Bell, S. D. and Jackson, S. P. (1997). Factor requirements for transcription in the Archaeon Sulfolobus shibatae. EMBO Journal 16, 2927-2936.

52. Singleton, B. K., Priestley, A., Steingrimsdottir, H., Gell, D., Blunt, T., Jackson, S. P., Lehmann, A. R. and Jeggo, P. A. (1997). Molecular and biochemical characterization of xrs mutants defective in Ku80. Mol Cell Biol 17, 1264–1273.

53. Teo, S.-H. and Jackson, S. P. (1997). Identification of Saccharomyces cerevisiae DNA ligase IV: involvement in DNA double-strand break repair. EMBO Journal 16, 4788- 4795.

54. Barlow, C., Liyanage, M., Moens, P. B., Tarsounas, M., Nagashima, K., Brown, K., Rottinghaus, S., Jackson, S. P., Tagle, D., Ried, T. and Wynshaw-Boris, A. (1998). Atm deficiency results in severe meiotic disruption as early as leptonema of prophase I. Development 125, 4007-4017.

55. Bell, S. D., Jaxel, C., Nadal, M., Kosa, P. F. and Jackson, S. P. (1998). Temperature, template topology, and factor requirements of archaeal transcription. Proc Natl Acad Sci USA 95, 15218-15222.

56. Boulton, S. J. and Jackson, S. P. (1998). Components of the Ku-dependent non- homologous end-joining pathway are involved in telomeric length maintenance and telomeric silencing. EMBO Journal 17, 1819-1828.

57. Errami, A., Finnie, N. J., Morolli, B., Jackson, S. P., Lohmann, P. H. M. and Zdzienicka, M. Z. (1998). Molecular and biochemical characterization of new X-ray-sensitive hamster cell mutants defective in Ku80. Nucleic Acids Research 26, 4332–4338.

58. Errami, A., He, D. M., Friedl, A. A., Overkamp, W. J. I., Morolli, B., Hendrickson, E. A., Eckardt-Schupp, F., Oshimura, M., Lohman, P. H. M., Jackson, S. P. and Zdzienicka, M. Z. (1998). XR-C1, a new CHO cell mutant which is defective in DNA-PKcs, is impaired in both V(D)J coding and signal joint formation. Nucleic Acids Research 26, 3146–3153.

59. Escarceller, M., Buchwald, M., Singleton, B. K., Jeggo, P. A., Jackson, S. P., Moustacchi, E. and Papadopoulo, D. (1998). Fanconi anemia C gene product plays a role in the fidelity of blunt DNA end-joining. Journal of Molecular Biology 279, 375- 385.

60. Freire, R., Murguia, J. R., Tarsounas, M., Lowndes, N. F., Moens, P. B. and Jackson, S. P. (1998). Human and mouse homologs of Schizosaccharomyces pombe rad1+ and Saccharomyces cerevisiae RAD17: linkage to checkpoint control and mammalian meiosis. Genes and Development 12, 2560-2573.

61. Priestley, A., Beamish, H. J., Gell, D., Amatucci, A. G., Muhlmann-Diaz, M. C., Singleton, B. K., Smith, G. C. M., Blunt, T., Schalkwyk, L. C., Bedford, J. S., Jackson, S. P., Jeggo, P. A. and Taccioli, G. E. (1998). Molecular and biochemical characterisation of DNA-dependent protein kinase-defective rodent mutant irs-20. Nucleic Acids Research 26, 1965-1973.

62. Qureshi, S. A. and Jackson, S. P. (1998). Sequence-specific DNA binding by the S. shibatae TFIIB homolog, TFB, and its effect on promoter strength. Molecular Cell 1, 389-400.

63. Taccioli, G. E., Amatucci, A. G., Beamish, H. J., Gell, D., Xiang, X. H., Arzayus, M. I. T., Priestley, A., Jackson, S. P., Rothstein, A. M., Jeggo, P. A. and Herrera, V. L. M. (1998). Targeted disruption of the catalytic subunit of the DNA-PK gene in mice confers severe combined immunodeficiency and radiosensitivity. Immunity 9, 355-366.

64. Trigon, S., Serizawa, H., Conaway, J. W., Conaway, R. C., Jackson, S. P. and Morange, M. (1998). Characterization of the residues phosphorylated in vitro by different C-terminal domain kinases. Journal of Biological Chemistry 273, 6769-6775.

65. Yavuzer, U., Smith, G. C. M., Bliss, T., Werner, D. and Jackson, S. P. (1998). DNA end- independent activation of DNA-PK mediated via association with the DNA-binding protein C1D. Genes and Development 12, 2188-2199.

66. Bell, S. D., Cairns, S. S., Robson, R. L. and Jackson, S. P. (1999). Transcriptional regulation of an archaeal operon in vivo and in vitro. Molecular Cell 4, 971-982.

67. Bell, S. D., Kosa, P. L., Sigler, P. B. and Jackson, S. P. (1999). Orientation of the transcription preinitiation complex in Archaea. Proc Natl Acad Sci USA 96, 13662- 13667.

68. d'Adda di Fagagna, F., Hande, M. P., Tong, W.-M., Lansdorp, P. M., Wang, Z.-Q. and Jackson, S. P. (1999). Functions of poly(ADP-ribose) polymerase in controlling telomere length and chromosomal stability. Nature Genetics 23, 76-80.

69. Downs, J. A. and Jackson, S. P. (1999). Involvement of DNA end-binding protein Ku in Ty element retrotransposition. Mol Cell Biol 19, 6260-6268.

70. Durocher, D., Henckel, J., Fersht, A. R. and Jackson, S. P. (1999). The FHA domain is a modular phosphopeptide recognition motif. Molecular Cell 4, 387-394.

71. Gell, D. and Jackson, S. P. (1999). Mapping of protein–protein interactions within the DNA-dependent protein kinase complex. Nucleic Acids Research 27, 3494-3502.

72. Izzard, R. A., Jackson, S. P. and Smith, G. C. M. (1999). Competitive and noncompetitive inhibition of the DNA-dependent protein kinase. Cancer Research 59, 2581-2586.

73. Lakin, N. D., Hann, B. C. and Jackson, S. P. (1999). The ataxia-telangiectasia related protein ATR mediates DNA-dependent phosphorylation of p53. Oncogene 18, 3989- 3995.

74. McAinsh, A. D., Scott-Drew, S., Murray, J. A. H. and Jackson, S. P. (1999). DNA damage triggers disruption of telomeric silencing and Mec1p-dependent relocation of Sir3p. Current Biology 9, 963-966.

75. Moens, P. B., Tarsounas, M., Morita, T., Habu, T., Rottinghaus, S. T., Freire, R., Jackson, S. P., Barlow, C. and Wynshaw-Boris, A. (1999). The association of ATR protein with mouse meiotic chromosome cores. Chromosoma 108, 95-102.

76. Plumb, M. A., Smith, G. C. M., Cunniffe, S. M. T., Jackson, S. P. and O'Neill, P. (1999). DNA-PK activation by ionizing radiation-induced DNA single-strand breaks. International Journal of Radiation Biology 75, 553-561.

77. Riballo, E., Critchlow, S. E., Teo, S.-H., Doherty, A. J., Priestley, A., Broughton, B., Kysela, B., Beamish, H., Plowman, N., Arlett, C. F., Lehmann, A. R., *Jackson, S. P. and *Jeggo, P. A. (1999). Identification of a defect in DNA ligase IV in a radiosensitive leukaemia patient. Current Biology 9, 699-702. * Co-corresponding authors.

78. Smith, G. C. M., d'Adda di Fagagna, F., Lakin, N. D. and Jackson, S. P. (1999). Cleavage and inactivation of ATM during apoptosis. Mol Cell Biol 19, 6076-6084.

79. Smith, G. C. M., Cary, R. B., Lakin, N. D., Hann, B. C., Teo, S.-H., Chen, D. J. and Jackson, S. P. (1999). Purification and DNA binding properties of the ataxia- telangiectasia gene product ATM. Proc Natl Acad Sci USA 96, 11134-11139.

80. Bell, S. D. and Jackson, S. P. (2000). The role of transcription factor B in transcription initiation and promoter clearance in the archaeon Sulfolobus acidocaldarius. Journal of Biological Chemistry 275, 12934-12940.

81. Bell, S. D. and Jackson, S. P. (2000). Mechanism of autoregulation by an archaeal transcriptional repressor. Journal of Biological Chemistry 275, 31624-31629.

82. Carlomagno, F., Burnet, N. G., Turesson, I., Nyman, J., Peacock, J. H., Dunning, A. M., Ponder, B. A. J. and Jackson, S. P. (2000). Comparison of DNA repair protein expression and activities between human fibroblast cell lines with different radiosensitivities. International Journal of Cancer 85, 845-849.

83. Downs, J. A., Lowndes, N. F. and Jackson, S. P. (2000). A role for Saccharomyces cerevisiae histone H2A in DNA repair. Nature 408, 1001-1004.

84. Durocher, D., Taylor, I. A., Sarbassova, D., Haire, L. F., Westcott, S. L., *Jackson, S. P., *Smerdon, S. J. and *Yaffe, M. B. (2000). The molecular basis of FHA domain : phosphopeptide binding specificity and implications for phospho-dependent signaling mechanisms. Molecular Cell 6, 1169-1182. * Co-corresponding authors.

85. Errami, A., Overkamp, W. J. I., He, D. M., Friedl, A. A., Gell, D. A., Eckardt-Schupp, F., Jackson, S. P., Hendrickson, E. A., Lohman, P. H. M. and Zdzienicka, M. Z. (2000). A new X-ray sensitive CHO cell mutant of ionizing radiation group 7, XR-C2, that is defective in DSB repair but has only a mild defect in V(D)J recombination. Mutation Research-DNA Repair 461, 59-69.

86. Moens, P. B., Freire, R., Tarsounas, M., Spyropoulos, B. and Jackson, S. P. (2000). Expression and nuclear localization of BLM, a chromosome stability protein mutated in Bloom's syndrome, suggest a role in recombination during meiotic prophase. Journal of Cell Science 113, 663-672.

87. Rouse, J. and Jackson, S. P. (2000). LCD1: an essential gene involved in checkpoint control and regulation of the MEC1 signalling pathway in Saccharomyces cerevisiae. EMBO Journal 19, 5801-5812.

88. Teo, S.-H. and Jackson, S. P. (2000). Lif1p targets the DNA ligase Lig4p to sites of DNA double-strand breaks. Current Biology 10, 165-168.

89. Wilson, C. R., Davidson, S. E., Margison, G. P., Jackson, S. P., Hendry, J. H. and West, C. M. L. (2000). Expression of Ku70 correlates with survival in carcinoma of the cervix. British Journal of Cancer 83, 1702-1706.

90. Bell, S. D., Brinkman, A. B., van der Oost, J. and Jackson, S. P. (2001). The archaeal TFIIE alpha homologue facilitates transcription initiation by enhancing TATA-box recognition. EMBO Reports 2, 133-138.

91. d'Adda di Fagagna, F., Hande, M. P., Tong, W.-M., Roth, D., Lansdorp, P. M., Wang, Z.-Q. and Jackson, S. P. (2001). Effects of DNA nonhomologous end-joining factors on telomere length and chromosomal stability in mammalian cells. Current Biology 11, 1192-1196.

92. D'Amours, D. and Jackson, S. P. (2001). The yeast Xrs2 complex functions in S phase checkpoint regulation. Genes and Development 15, 2238-2249.

93. Doherty, A. J., Jackson, S. P. and Weller, G. R. (2001). Identification of bacterial homologues of the Ku DNA repair proteins. FEBS Letters 500, 186-188.

94. Eichhorn, K. and Jackson, S. P. (2001). A role for TAF3B2 in the repression of human RNA polymerase III transcription in nonproliferating cells. Journal of Biological Chemistry 276, 21158-21165.

95. Freire, R., di Fagagna, F. D., Wu, L., Pedrazzi, G., Stagljar, I., Hickson, I. D. and Jackson, S. P. (2001). Cleavage of the Bloom's syndrome gene product during apoptosis by caspase-3 results in an impaired interaction with topoisomerase III a. Nucleic Acids Research 29, 3172-3180.

96. Herceg, Z., Hulla, W., Gell, D., Cuenin, C., Lleonart, M., Jackson, S. and Wang, Z. Q. (2001). Disruption of Trrap causes early embryonic lethality and defects in cell cycle progression. Nature Genetics 29, 206-211.

97. Magill, C. P., Jackson, S. P. and Bell, S. D. (2001). Identification of a conserved archaeal RNA polymerase subunit contacted by the basal transcription factor TFB. Journal of Biological Chemistry 276, 46693-46696.

98. Sibanda, B. L., Critchlow, S. E., Begun, J., Pei, X. Y., Jackson, S. P., Blundell, T. L. and Pellegrini, L. (2001). Crystal structure of an Xrcc4–DNA ligase IV complex. Nature Structural Biology 8, 1015–1019.

99. Teo, S.-H. and Jackson, S. P. (2001). Telomerase subunit overexpression suppresses telomere-specific checkpoint activation in the yeast yku80 mutant. EMBO Reports 2, 197-202.

100. Bell, S. D., Botting, C. H., Wardleworth, B. N., Jackson, S. P. and White, M. F. (2002). The interaction of Alba, a conserved archaeal, chromatin protein, with Sir2 and its regulation by acetylation. Science 296, 148-151.

101. Chang, J.-H., Kim, H.-C., Hwang, K.-Y., Lee, J.-W., Jackson, S. P., Bell, S. D. and Cho, Y. (2002). Structural basis for the NAD-dependent deacetylase mechanism of Sir2. Journal of Biological Chemistry 277, 34489-34498.

102. Li, J., Williams, B. L., Haire, L. F., Goldberg, M., Walker, E., Durocher, D., Yaffe, M. B., *Jackson, S. P. and *Smerdon, S. J. (2002). Structural and functional versatility of the FHA domain in DNA-damage signaling by the tumor suppressor kinase Chk2. Molecular Cell 9, 1045-1054. * Co-corresponding authors.

103. Rouse, J. and Jackson, S. P. (2002). Lcd1p recruits Mec1p to DNA lesions in vitro and in vivo. Molecular Cell 9, 857-869.

104. Weller, G. R., Kysela, B., Roy, R., Tonkin, L. M., Scanlan, E., Della, M., Devine, S. K., Day, J. P., Wilkinson, A., d'Adda di Fagagna, F., Devine, K. M., Bowater, R. P., Jeggo, P. A., Jackson, S. P. and Doherty, A. J. (2002). Identification of a DNA nonhomologous end-joining complex in bacteria. Science 297, 1686-1689.

105. d'Adda di Fagagna, F., Weller, G. R., Doherty, A. J. and Jackson, S. P. (2003). The Gam protein of bacteriophage Mu is an orthologue of eukaryotic Ku. EMBO Reports 4, 47-52.

106. d'Adda di Fagagna, F., Reaper, P. M., Clay-Farrace, L., Fiegler, H., Carr, P., Von Zglinicki, T., Saretzki, G., Carter, N. P. and Jackson, S. P. (2003). A DNA damage checkpoint response in telomere-initiated senescence. Nature 426, 194-198.

107. Dionne, I., Nookala, R. K., Jackson, S. P., Doherty, A. J. and Bell, S. D. (2003). A heterotrimeric PCNA in the hyperthermophilic archaeon Sulfolobus solfataricus. Molecular Cell 11, 275-282.

108. Downs, J. A., Kosmidou, E., Morgan, A. and Jackson, S. P. (2003). Suppression of homologous recombination by the Saccharomyces cerevisiae linker histone. Molecular Cell 11, 1685-1692.

109. Goldberg, M., Stucki, M., Falck, J., D'Amours, D., Rahman, D., Pappin, D., Bartek, J. and Jackson, S. P. (2003). MDC1 is required for the intra-S-phase DNA damage checkpoint. Nature 421, 952-956.

110. Downs, J. A., Allard, S., Jobin-Robitaille, O., Javaheri, A., Auger, A., Bouchard, N., Kron, S. J., Jackson, S. P. and Cote, J. (2004). Binding of Chromatin-Modifying Activities to Phosphorylated Histone H2A at DNA Damage Sites. Molecular Cell 16, 979-990.

111. Hickson, I., Zhao, Y., Richardson, C. J., Green, S. J., Martin, N. M., Orr, A. I., Reaper, P. M., Jackson, S. P., Curtin, N. J. and Smith, G. C. (2004). Identification and characterization of a novel and specific inhibitor of the ataxia-telangiectasia mutated kinase ATM. Cancer Research 64, 9152-9159.

112. Jazayeri, A., McAinsh, A. D. and Jackson, S. P. (2004). Saccharomyces cerevisiae Sin3p facilitates DNA double-strand break repair. Proc Natl Acad Sci U S A 101, 1644- 1649.

113. Lau, A., Kanaar, R., Jackson, S. P. and O'Connor, M. J. (2004). Suppression of retroviral infection by the RAD52 DNA repair protein. EMBO Journal 23, 3421-3429.

114. Lukas, C., Melander, F., Stucki, M., Falck, J., Bekker-Jensen, S., Goldberg, M., Lerenthal, Y., Jackson, S. P., Bartek, J. and Lukas, J. (2004). Mdc1 couples DNA double-strand break recognition by Nbs1 with its H2AX-dependent chromatin retention. EMBO Journal 23, 2674-2683.

115. Riballo, E., Kuhne, M., Rief, N., Doherty, A., Smith, G. C., Recio, M. J., Reis, C., Dahm, K., Fricke, A., Krempler, A., Parker, A. R., Jackson, S. P., Gennery, A., Jeggo, P. A. and Lobrich, M. (2004). A Pathway of Double-Strand Break Rejoining Dependent upon ATM, Artemis, and Proteins Locating to gamma-H2AX Foci. Molecular Cell 16, 715- 724.

116. Roy, R., Meier, B., McAinsh, A. D., Feldmann, H. M. and Jackson, S. P. (2004). Separation-of-function mutants of yeast Ku80 reveal a Yku80p-Sir4p interaction involved in telomeric silencing. J Biol Chem 279, 86-94.

117. Squires, S., Coates, J. A., Goldberg, M., Toji, L. H., Jackson, S. P., Clarke, D. J. and Johnson, R. T. (2004). p53 Prevents the Accumulation of Double-Strand DNA Breaks at Stalled-Replication Forks Induced by UV in Human Cells. Cell Cycle 3, 1543-1557.

118. Falck, J., Coates, J. and Jackson, S. P. (2005). Conserved modes of recruitment of ATM, ATR and DNA-PKcs to sites of DNA damage. Nature 434, 605-611.

119. Giavara, S., Kosmidou, E., Hande, M. P., Bianchi, M. E., Morgan, A., d'Adda di Fagagna, F. and Jackson, S. P. (2005). Yeast Nhp6A/B and Mammalian Hmgb1 Facilitate the Maintenance of Genome Stability. Current Biology 15, 68-72.

120. Harvey, A. C., Jackson, S. P. and Downs, J. A. (2005). Saccharomyces cerevisiae Histone H2A Ser122 facilitates DNA repair. Genetics 170, 543-553.

121. Lee, A. C., Fernandez-Capetillo, O., Pisupati, V., *Jackson, S. P. and Nussenzweig, A. (2005). Specific Association of Mouse MDC1/NFBD1 with NBS1 at Sites of DNA- Damage. Cell Cycle 4, 177-182.

122. Farmer, H., McCabe, N., Lord, C. J., Tutt, A. N. J., Johnson, D. A., Richardson, T. B., Santorosa, M., Dillon, K. J., Hickson, I., Martin, N. M. B., Jackson, S. P., Smith, G. C. M. and Ashworth, A. (2005). Targeting the DNA repair defect in BRCA mutant cells as a therapeutic strategy. Nature 434, 917-921.

123. Lau, A., Swinbank, K.M., Ahmed, P.S., Taylor, D.L., Jackson, S.P., Smith, G.C.M. and O'Connor, M.J. (2005). Suppression of HIV-1 infection by a small molecule inhibitor of the ATM kinase. Nat Cell Biol. 7, 493-500.

124. Tutt, A.N.J., Lord, C.J., McCabe, N., Farmer, H., Turner, N., Martin, N.M., Jackson, S.P., Smith, G.C.M. and Ashworth, A. (2005). Exploiting the DNA repair defect in BRCA mutant cells in the design of new therapeutic strategies for cancer. Cold Spring Harbor Symposia on Quantitative Biology, 139-148.

125. Moumen, A., Masterson, P., O'Connor, M. and Jackson, S.P. (2005). hnRNP K: An HDM2 target and transcriptional co-activator of p53 in response to DNA damage. Cell 123, 1065-1078

126. Stucki, M., Clapperton, J.A, Mohammad, D., Yaffe, M.B., Smerdon, S.J. and Jackson, S.P. (2005). MDC1 directly binds phosphorylated histone H2AX to regulate cellular responses to DNA double-strand breaks. Cell 123, 1213-1226

127. Short, S.C., Bourne, S., Martindale, C., Woodcock, M. and Jackson, S.P. (2005). DNA damage responses at low radiation doses. Radiation Research 164, 292-302

128. Jazayeri, A., Falck, J., Lukas, C., Bartek, J., Lukas, J., Smith, G.C.M. and Jackson, S.P. (2006). ATM- and cell cycle-dependent regulation of ATR in response to DNA double-strand breaks. Nat Cell Biol. 8, 37-45

129. Smits, V.A.J., Reaper, P.M. and Jackson, S.P. (2006). Rapid PIKK-dependent release of Chk1 from chromatin promotes the DNA-damage checkpoint response. Current Biology 16, 150-159.

130. Grenon, M., Magill, C., Lowndes, N. and Jackson, S.P. (2006). Double-strand breaks trigger MRX- and Mec1-dependent, but Tel1 independent, checkpoint activation. FEMS Yeast Research 1-12.

131. Ahnesorg, P., Smith, P. and Jackson, S.P. (2006). XLF interacts with the XRCC4-DNA ligase IV complex to promote DNA non-homologous end-joining. Cell 124, 301-313

132. Henderson, M.J., Munoz, M.A., Saunders, D.N., Clancy, J.L., Russell, A.J., Williams, B., Pappin, D. Khanna, K.K., Jackson, S.P., Sutherland, R.L. and Watts, C.K.W. (2006). EDD mediate DNA damage-induced activation of CHK2. J Biol Chem 281, 39990- 40000

133. Dore A.S., Furnham N., Davies O.R., Sibanda B.L., Chirgadze D.Y., Jackson S.P., Pellegrini L. and Blundell T.L. (2006). Structure of an Xrcc4-DNA ligase IV yeast ortholog complex reveals a novel BRCT interaction mode. DNA Repair (Amst). 5(3), 362-368

134. Ahnesorg, P. and Jackson, S.P. (2007). The non-homologous end-joining protein Nej1p is a target of the DNA damage checkpoint. DNA Repair 6,190-201.

135. Hentges, P., Ahnesorg, P., Pitcher, R.S., Bruce, C.K., Kysela, B., Green, A.J., Bianchi, J., Wilson, T.E., Jackson, S.P. and Doherty, A.J. (2007). Evolutionary and functional conservation of the DNA non homologous end joining protein, XLF/Cernunnos. J Biol Chem 281, 37517-37526

136. Driscoll, R. Hudson, A. and Jackson, S.P. (2007). Rtt109 promotes genome stability by acting as an acetyl-transferase for histone H3 lysine 56. Science 315, 649-652.

137. Meier A., Fiegler, H., Muñoz P., Rigler D., Langford C., Blasco M., Carter N. and Jackson S.P. (2007). Spreading of mammalian DDR factors studied by ChIP-chip at damaged telomeres. EMBO J. 26, 2707-2718

138. Xhemalce, B., Miller, K.M., Driscoll, R., Masumoto, H., Jackson, S.P., Kouzarides, T., Verreault, A. and Arcangioli, B. (2007). Regulation of histone H3 lysine 56 acetylation in Schizosaccharomyces pombe. J Biol Chem. 282, 15040-15047

139. Bartkova, J., Horejsi, Z., Sehested, M., Nesland, J.M., Rajpert-De Meyts, E., Skakkebaek, N.E., Stucki, M., Jackson, S.P., Lukas, J. and Bartek, J. (2007). DNA damage response mediators MBD1 and 53BP1: Constitutive activation and aberrant loss in breast and lung cancer, but not in testicular germ cell tumours. Oncogene 26, 7414-7422

140. Difilippantonio S., Celeste A., Kruhlak M.J., Lee Y., Difilippantonio M.J., Feigenbaum L., Jackson S.P., McKinnon P.J., and Nussenzweig A. (2007) Distinct domains in Nbs1 regulate irradiation-induced checkpoints and apoptosis. J Exp Med. 204, 1003-1011

141. Hammet, A., Magill, C., Heierhorst, J. and Jackson, S.P. (2007) Rad9 BRCT domain interaction with phosphorylated H2AX regulates the G1 checkpoint in budding yeast. EMBO Reports. 8, 851-857

142. Sartori, A. A., Lukas, C., Coates, J., Mistrik, M., Fu, S., Bartek, J., Baer, R., Lukas, J., and Jackson, S. P. (2007). Human CtIP promotes DNA end resection. Nature 450, 509- 514.

143. Li, Y., Chirgadze, D.Y., Bolanos-Garcia, V.M., Sibanda, B.L., Davies, O.R., Ahnesorg, P., Jackson, S.P. and Blundell, T.L. (2007). Crystal structure of human XLF/Cernunnos reveals unexpected differences from XRCC4 with implications for NHEJ. EMBO J. 27, 290-300.

144. Kolas, N.K., Chapman, J.R., Nakada, S., Ylanko, J., Chahwan, R., Sweeney, F.D., Panier, S., Mendez, M., Wildenhain, J., Thomson, T.M., Pelletier, L., Jackson, S.P*. and Durocher, D*. (2007). Orchestration of the DNA-damage response by the RNF8 ubiquitin ligase. Science 318, 1637-1640. * Co-corresponding authors.

145. Xie A., Hartlerode A., Stucki M., Odate S., Puget N., Kwok A., Nagaraju G., Yan C., Alt F.W., Chen J., Jackson S.P. and Scully R. (2007). Distinct Roles of Chromatin- Associated Proteins MDC1 and 53BP1 in Mammalian Double-Strand Break Repair. Mol Cell. 28(6), 1045-1057.

146. Chapman J.R. and Jackson S.P. (2008), Phospho-dependent interactions between NBS1 and MDC1 mediate chromatin retention of the MRN complex at sites of DNA damage. EMBO Reports 9, 795-801.

147. Huertas P., Cortes-Ledesma F., Sartori, A.A., Aguilera, A. and Jackson, S.P. (2008). CDK targets Sae2 to control DNA-end resection and homologous recombination. Nature 455, 689-692.

148. Gravel, S., Chapman, J.R., Magill, C. and Jackson, S.P. (2008). DNA helicases Sgs1 and BLM promote DNA double-strand break resection. Genes and Development 22, 2767-2772.

149. Enge, M., Bao, W., Hedstrom, E., Jackson, S.P., Moumen, A. and Selivanova, G. (2009). MDM2-dependent downregulation of p21 and hnRNPK provides a switch between apoptosis and growth arrest induced by pharmacologically activated p53. Cancer Cell 15, 171-183.

150. Sollier, J., Driscoll, R., Castellucci, F., Foiani, M., Jackson, S.P. and Branzei D. (2009). The Saccharomyces cerevisiae Esc2 and Smc5-6 proteins promote sister chromatid junction-mediated intra-S repair. Molecular Biology of the Cell 20, 1671-1682.

151. Huertas, P. and Jackson, S.P. (2009). Human CtIP mediates cell cycle control of DNA end resection and double strand break repair. Journal of Biological Chemistry, 284, 9558-9565.

152. Tjeertes, J.V., Miller, K.M. and Jackson, S.P. (2009). Screen for DNA-damage- responsive histone modifications identifies H3K9Ac and H3K56Ac in human cells. The EMBO Journal 28, 1878-1889.

153. Ahel, D., Hofejsi, Z., Wiechens, N., Polo, S.E., Garcia-Wilson, E., Ahel, I., Flynn, H., Skehel, M., West, S.C., Jackson, S.P., Owen-Hughes, T. and Boulton, S.J. (2009). Poly (ADP-ribose)-dependent regulation of DNA repair by the chromatin remodeling enzyme ALC1. Science, 325, 1240-1243.

154. Lloyd, J., Chapman, J.R., Clapperton, J.A., Haire, L.F., Hartsuiker, E., Li, J., Carr, A.M., *Jackson, S.P. and *Smerdon, S.J. (2009). A supra-modular FHA/BRCT-repeat architecture mediates Nbs1 adaptor function in response to DNA-damage. Cell. 139, 100-111. * Co-corresponding authors.

155. Galanty, Y., Belotserkovskaya, R., Coates, J., Polo, S., Miller, K.M. and Jackson, S.P. (2009). Mammalian SUMO E3-ligases PIAS1 and PIAS4 promote responses to DNA double-strand breaks. Nature 462, 935-939.

156. Polo, S.E., Kaidi, A., Baskcomb, L., Galanty, Y. and Jackson, S.P. (2010). Regulation of DNA damage responses and cell cycle progression by the chromatin remodeling factor CHD4. EMBO Journal 29, 3130-3139.

157. Miller, K.M, Tjeertes, J.V., Coates, J., Legube, G., Polo, S.E., Britton, S. and Jackson, S.P. (2010). Human HDAC1 and HDAC2 function in the DNA-damage response to promote DNA non-homologous end-joining. Nature Structural and Molecular Biology 17, 1144-1151.

158. Hajkova, P., Jeffries, S.J., Lee, C., Miller, N., Jackson, S.P. and Surani, M.A. (2010). Genome-wide reprogramming in the mouse germ line entails the base excision repair pathway. Science 329, 78-82.

159. Giunta, S., Belotserkovskaya, R. and Jackson, S.P. (2010). DNA damage signaling in response to double-strand breaks during mitosis. Journal of Cell Biology 190, 197-207.

160. Kaidi, A., Weinert, B. T., Choudhary, C., and Jackson, S. P. (2010). Human SIRT6 promotes DNA end resection through CtIP deacetylation. Science 329, 1348-1353.

161. Harrigan, J. A., Belotserkovskaya, R., Coates, J., Dimitrova, D. S., Polo, S. E., Bradshaw, C. R., Fraser, P., and Jackson, S. P. (2011). Replication stress induces 53BP1- containing OPT domains in G1 cells. Journal of Cell Biology, 193, 97-108.

162. Flott, S., Kwon, Y., Pigli, Y.Z., Rice, P.A., Sung, P. and Jackson, S.P. (2011). Regulation of Rad51 function by phosphorylation. EMBO Reports 12(8), 833-9.

163. Blasius, M., Forment, J.V., Thakker, N., Wagner, S.A., Choudhary, C. and Jackson, S.P. (2011). A phospho-proteomic screen identifies substrates of the checkpoint kinase Chk1. Genome Biology, 12(8):R78.

164. Qvist, P., Huertas, P.*, Jimeno, S., Nyegaard, M., Hassan, M.J., Jackson, S.P.* and Børglum, A.D.* (2011). CtIP mutations cause Seckel and Jawad syndromes. PLoS Genetics 7(10): e1002310. *Co-corresponding authors.

165. Forment J.V., Blasius M., Guerini I., Jackson S.P. (2011) Structure-Specific DNA Endonuclease Mus81/Eme1 Generates DNA Damage Caused by Chk1 Inactivation. PLoS ONE 6(8): e23517.

166. Rodriguez, R., Miller, K.M., Forment, J.V., Bradshaw, C.R., Mehran, N., Britton, S., Oelschlagel, T., Xhemalce, B., Balasubramanian, S*., and Jackson, S.P*. (2012). Small molecule-induced DNA damage identifies alternative DNA structures in human genes. Nature Chemical Biology, 8, 301-310. *Co-corresponding authors.

167. Polo, S.E., Blackford, A.N., Chapman, J.R., Baskcomb, L., Gravel, S., Rusch, A., Thomas, A., Blundred, R., Smith, P., Kzhyshkowska, J., Dobner, T., Taylor, A.M.R., Turnell, A.S., Stewart, G.S. Grand, R.J* and Jackson, S.P*. (2012) Regulation of DNA- end resection by hnRNPU-like proteins promotes DNA double-strand break signaling and repair. Molecular Cell 45, 505-516. *Co-corresponding authors

168. Beli, P., Lukashchuk, N., Wagner, S.A., Weinert, B.T., Olsen, J.V., Baskcomb, L., Mann, M., Jackson, S.P*. and Choudhary, C*. (2012) Proteomic investigations reveal a role for RNA processing factor THRAP3 in the DNA damage response. Molecular Cell 46, 212-25. *Co-corresponding authors.

169. Chapman, J.R., Sossick, A., Boulton, S.J. and Jackson, S.P. (2012) BRCA1-associated exclusion of 53BP1 from DNA damage sites underlies temporal control of DNA repair. Journal of Cell Science 125, 3529-3534.

170. Schiller, C.B., Lammens, K., Guerini, I., Coordes, B., Feldmann, H., Schlauderer, F., Möckel, C., Schele, A., Sträßer, K., Jackson, S.P. and Hopfner, K-P. (2012) Structural biology of the Mre11:Nbs1 complex yields insights into ataxia-telangiestasia-like disease mutations and DNA damage signaling. Nature Structural and Molecular Biology 19, 693-700.

171. Falck, J., Forment, J.V., Coates, J., Mistrik, M., Lukas, J., Bartek, J. and Jackson, S.P. (2012) CDK targeting of NBS1 promotes DNA-end resection, replication restart and homologous recominbation. EMBO Reports, 13, 561-568.

172. Galanty, Y., Belotserkovskaya, R., Coates, J., and Jackson, S. P. (2012). RNF4, a SUMO-targeted ubiquitin E3 ligase, promotes DNA double-strand break repair. Genes and Development 26, 1179-1195.

173. Forment, J.V, Walker, R.V and Jackson, S.P. (2012). A high-throughput, flow cytometry-based method to quantify DNA-end resection in mammalian cells. Cytometry 81A, 922-928.

174. McIntyre, R.E., Lakshminarasimhan, P., Ismail, O., Carragher, D.M., Sanchez-Andrade, G., Forment, J.V., Fu, B., Del Castillo Velasco Herrera, M., Edwards, A., Yang, F., Sanger Mouse Genetics Project, Ramirez-Solis, R., Estabel, J., Gallagher, F.A., Logan, D.W., Arends, M.J., Tsang, S.H., Mahajan, V.B., Scudamore, C.L., White, J.K., Jackson, S.P., Gergely, F., Adams, D.J. (2012). Disruption of Mouse Cenpj, a Regulator of Centriole Biogenesis, Phenocopies Seckel Syndrome. PLoS Genetics, 8

175. Chen, W-T., Alpert, A., LeiterC., Gonh, F., Jackson, S.P*. and Miller, K*. (2013). Systematic identification of functional residues in mammalian histone H2AX. Molecular and Cellular Biology 33, 111-26 *Co-corresponding authors.

176. Munoz-Galvan, S., Lopez-Saavedra, A., Jackson, S.P., Huertas, P., Cortes-Ledesma, F. and Aguilera, A. (2013). Competing roles of DNA end resection and non- homologous end joining functions in the repair of replication-born double-strand breaks by sister-chromatid recombination. Nucleic Acids Research 41, 1669-1683

177. Moumen, A*., Magill, C., Dry, K.L. and Jackson, S.P*. (2013). ATM-dependent phosphorylation of heterogeneous nuclear ribonucleoprotein K promotes p53 transcriptional activation in response to DNA damage. Cell Cycle 12, 698-704. *Co- corresponding authors.

178. Chahwan, R., Gravel, S., Matsusaka, T. and Jackson, S.P. (2013). Dma/RNF8 proteins are evolutionarily conserved E3 ubiquitin ligases that target septins. Cell Cycle, 12, 1000-1008.

179. Kaidi, A. and Jackson, S.P. (2013). KAT5 tyrosine phosphorylation couples chromatin sensing to ATM checkpoint signalling. Nature 498, 70-74.

180. Britton, S., Coates, J. and Jackson, S.P. (2013). A new method for high-resolution imaging of Ku foci to decipher mechanisms of DNA double-strand break repair. Journal of Cell Biology 202, 579-595.

181. Larrieu, D., Britton, S., Demir, M., Rodriguez, R*. and Jackson, S.P*. (2014). Chemical inhibition of NAT10 corrects defects of laminopathic cells. Science 344, 527-532. *Co- corresponding authors.

182. Xi, W., Schmidt, C.K., Sanchez, S., Gracias, D.H., Carozo-Salas, R.E., Jackson, S.P. and Schmidt, O.G. (2014) Rolled-up functionalized nanomembranes as three- dimensional cavities for single cell studies. NANOLetters 14, 4197-4204.

183. Aymard, F., Bugler, B., Schmidt, C.K., Guillou, E., Caron, P., Briois, S., Iacovoni, J.S., Daburon, V., Miller, K.M., Jackson, S.P. and Legube, G. (2014). Transcriptionally active chromatin recruits homologous recombination at DNA double-strand breaks. Nature Structural and Molecular Biology 21, 366-374.

184. Knobel, P.A., Belotserkovskaya, R., Galanty, Y., Schmidt, C.K., Jackson, S.P*. and Stracker, T.H*. (2014). USP28 is recruited to sites of DNA damage by the tansdem BRCT domains of 53BP1 but plays a minor role in double-strand break metabolism. Molecular and Cellular Biology 34, 2062-2074. *Co-corresponding authors.

185. Koch, B., Sanchez, S., Schmidt, C.K., Swiersy, A., Jackson, S.P., Schmidt, O.G. (2014). Confinement and deformation of single cells and their nuclei inside size-adapted microtubes. Advanced Healthcare Materials April 25 [Epub ahead of print].

186. Polato, F., Callen, E., Wong, N., Faryabi, R., Bunting, S., Chen, H.T., Kozak, M., Kruhlak, M.J., Reczek, C.R., Lee, W.H., Ludwig, T., Baer, R., Feigenbaum, L., Jackson, S., Nussenzweig, A. (2014). CtIP-mediated resection is essential for viability and can operate independently of BRCA1. Journal of Experimental Medicine 211, 1027-1036.

187. Blasius, M., Wagner, S.A., Choudhary, C., Bartek, J., Jackson, S.P. (2014). A quantitative 14-3-3 interaction screen connects the nuclear exosome targeting complex to the DNA damage response. Genes and Development 28, 1977-1982.

188. Nishi, R., Wijnhoven, P., le Sage, C., Tjeertes, J., Galanty, Y., Forment, J.V., Clague, M.J., Urbé, S., Jackson, S.P. (2014). Systematic characterization of deubiquitylating enzymes for roles in maintaining genome integrity. Nature Cell Biology 16, 1016-1026.

189. Davies O.R., Forment J.V., Sun M., Belotserkovskaya R., Coates J., Galanty Y., Demir M., Morton C.R., Rzechorzek N.J., Jackson S.P.*, Pellegrini L*. (2015) CtIP tetramer assembly is required for DNA-end resection and repair. Nature Structural and Molecular Biology Advance Online Publication. *Co-corresponding author.

190. Balmus G., Karp N.A., Ng B.L., Jackson S.P., Adams D.J., McIntyre R.E. (2015) A high- throughput in vivo micronucleus assay for genome instability screening in mice. Nature Protocols 10, 205-215.

191. Ochi T., Blackford A.N., Coates J., Jhujh S., Mehmood S., Tamura N., Travers J., Wu Q., Draviam V.M., Robinson C.V., Blundell T.L.*, Jackson S.P.* (2015) PAXX, a paralog of XRCC4 and XLF, interacts with Ku to promote DNA double-strand break repair. Science 347, 185-188. *Co-corresponding author.

192. Blackford, A.N., Nieminuszczy, J., Schwab, R.A., Galanty, Y., Jackson, S.P.*, and Niedzwiedz, W.* (2015). *Co-corresponding author. TopBP1 interacts with BLM to maintain genome stability but is dispensable for preventing BLM degradation. Molecular Cell 57, 1133-41.

193. Puddu, F., Oelschlaegel, T., Guerini, I., Geisler, N.J., Niu, H., Herzog, M., Salguero, I., Ochoa-Montano, B., Vire, E., Sung, P., Adams, D.J., Keene, T.M. and Jackson, S.P. (2015). Synthetic viability genomic screening defines Sae2 function in DNA repair. EMBO Journal 34, 1509-22

194. Brown JS, Lukashchuk N, Sczaniecka-Clift M, Britton S, le Sage C, Calsou P, Beli P, Galanty Y, Jackson SP. (2015) Neddylation Promotes Ubiquitylation and Release of Ku from DNA-Damage Sites. Cell Reports 11, 704-714.

195. Forment JF, Jackson SP (2015) A flow cytometry-based method to simplify the analysis and quantification of protein association to chromatin in mammalian cells. Nature Protocols 10, 1297-1307.

196. Yuan, Y., Britton, S., Delteil, C., Coates, J., Jackson, S.P., Barboule, N., Frit, P. and Calsou, P. (2015) Single-stranded DNA oligomers stimulate error-prone alternative repair of DNA double-strand breaks through hijacking Ku protein. Nucleic Acids Research Sep 8 (in press)

197. Wijnhoven, P., Konietzny, R., Blackford, A.N., Travers, J., Kessler, B.M., Nishi, R. and Jackson, S.P. (2015) USP4 promotes DNA repair and is subject to ubiquitylation autoregulation (in press)

198. Schmidt, C., Galanty, Y., Sczaniecka-Clift, M., Coates, J., Jhujh, S., Demir, M., Cornwell, M., Beli, P. and Jackson, S.P. (2015) Systematic E2 screening reveals a UBE2D-RNF138-CtIP axis promoting DNA repair. Nature Cell Biology, in press.

Review articles:

1. White, R. J. and Jackson, S. P. (1992). The TATA-binding protein: a central role in transcription by RNA polymerase I, polymerase II and polymerase III. Trends in Genetics 8, 284-288.

2. Jackson, S. P. (1992). Regulating transcription factor activity by phosphorylation. Trends in Cell Biology 2, 104-108.

3. Finnie, N., Gottlieb, T., Hartley, K. and Jackson, S. P. (1993). Transcription factor phosphorylation by the DNA-dependent protein kinase. Biochemical Society Transactions 21, 930-935.

4. Jackson, S., Gottlieb, T. and Hartley, K. (1993). Phosphorylation of transcription factor SP1 by the DNA-dependent protein kinase. Advances in Second Messenger and Phosphoprotein Research 28, 279-286.

5. Jackson, S.P., Identification and characterization of eukaryotic transcription factors, in Gene transcription: a practical approach, B.D. Hames and S.J. Higgins, Editors. 1993, IRL Press: Oxford. p. 189–242.

6. Gottlieb, T. M. and Jackson, S. P. (1994). Protein kinases and DNA damage. Trends in Biochemical Sciences 19, 500-503.

7. Jeggo, P. A., Taccioli, G. E. and Jackson, S. P. (1995). Menage à trois: double-strand break repair, V(D)J recombination and DNA-PK. BioEssays 17, 949-957.

8. Jackson, S. P. (1995). Ataxia–telangiectasia at the crossroads. Current Biology 5, 1210–1212.

9. Baumann, P., Qureshi, S. A. and Jackson, S. P. (1995). Transcription: new insights from studies on Archaea. Trends in Genetics 11, 279-283.

10. Jackson, S. P. and Jeggo, P. A. (1995). DNA double-strand break repair and V(D)J recombination: involvement of DNA-PK. Trends in Biochemical Sciences 20, 412-415.

11. Jackson, S. P. (1996). DNA damage detection by DNA-dependent protein kinase and related enzymes. Cancer Surveys 28, 261-279.

12. Finnie, N. J., Gottlieb, T. M., Blunt, T., Jeggo, P. A. and Jackson, S. P. (1996). DNA- dependent protein kinase defects are linked to deficiencies in DNA repair and V(D)J recombination. Philosophical Transactions of the Royal Society of London Series B- Biological Sciences 351, 173-179.

13. Jeggo, P. A., Jackson, S. P. and Taccioli, G. E. (1996). Identification of the catalytic subunit of DNA dependent protein kinase as the product of the mouse scid gene. Current Topics in Microbiology and Immunology 217, 79-89.

14. Jackson, S. P. (1996). The recognition of DNA damage. Current Opinion in Genetics and Development 6, 19-25.

15. Jackson, S. P. (1997). DNA-dependent protein kinase. International Journal of Biochemistry and Cell Biology 29, 935-938.

16. Jackson, S. P. (1997). Genomic stability: silencing and DNA repair connect. Nature 388, 829-830.

17. Elgin, S. C. R. and Jackson, S. P. (1997). Chromosomes and expression mechanisms. Current Opinion in Genetics & Development 7, 149-151.

18. Featherstone, C. and Jackson, S. P. (1998). DNA repair: the Nijmegen breakage syndrome protein. Current Biology 8, R622-R625.

19. Critchlow, S. E. and Jackson, S. P. (1998). DNA end joining: from yeast to man. Trends in Biochemical Sciences 23, 394-398.

20. Bell, S. D. and Jackson, S. P. (1998). Transcription in Archaea. Cold Spring Harbor Symposia on Quantitative Biology 63, 41-51.

21. Bell, S. D. and Jackson, S. P. (1998). Transcription and translation in Archaea: a mosaic of eukaryal and bacterial features. Trends in Microbiology 6, 222-228.

22. Smith, G. C. M. and Jackson, S. P. (1999). The DNA-dependent protein kinase. Genes and Development 13, 916-934.

23. Jackson, S. P. (1999). Detection, repair and signalling of DNA double-strand breaks. Biochemical Society Transactions 27, 1-13.

24. Smith, G. C. M., Divecha, N., Lakin, N. D. and Jackson, S. P. (1999). DNA-dependent protein kinase and related proteins. Biochemical Society Symposium, 91-104.

25. Featherstone, C. and Jackson, S. P. (1999). Ku, a DNA repair protein with multiple cellular functions? Mutation Research-DNA Repair 434, 3-15.

26. Featherstone, C. and Jackson, S. P. (1999). DNA-dependent protein kinase gets a break: its role in repairing DNA and maintaining genomic integrity. British Journal of Cancer 80, 14-19.

27. Lakin, N. D. and Jackson, S. P. (1999). Regulation of p53 in response to DNA damage. Oncogene 18, 7644-7655.

28. Featherstone, C. and Jackson, S. P. (1999). DNA double-strand break repair. Current Biology 9, R759-R761.

29. Bell, S. D. and Jackson, S. P. (2000). Charting a course through RNA polymerase. Nature Structural Biology 7, 703-705.

30. Durocher, D., Smerdon, S. J., Yaffe, M. B. and Jackson, S. P. (2000). The FHA domain in DNA repair and checkpoint signaling. Cold Spring Harbor Symposia on Quantitative Biology 65, 423-431.

31. McAinsh, A. D., Bell, S. D. and Jackson, S. P. (2000). Sir2: a key player in silencing, ageing and metabolism. The ELSO Gazette: e-magazine of the European Life Scientist Organization (http://www.the-elso-gazette.org).

32. Durocher, D. and Jackson, S. P. (2001). DNA-PK, ATM and ATR as sensors of DNA damage: variations on a theme? Current Opinion in Cell Biology 13, 225-231.

33. Khanna, K. K. and Jackson, S. P. (2001). DNA double-strand breaks: signaling, repair and the cancer connection. Nature Genetics 27, 247-254.

34. Bell, S. D. and Jackson, S. P. (2001). Mechanism and regulation of transcription in archaea. Current Opinion in Microbiology 4, 208-213.

35. Bell, S. D., Magill, C. P. and Jackson, S. P. (2001). Basal and regulated transcription in Archaea. Biochemical Society Transactions 29, 392-395.

36. Doherty, A. J. and Jackson, S. P. (2001). DNA repair: how Ku makes ends meet. Current Biology 11, R920-R924.

37. Jackson, S. P. (2001). Detecting, signalling and repairing DNA double-strand breaks. Biochemical Society Transactions 29, 655-661.

38. Khanna, K. K., Lavin, M. F., Jackson, S. P. and Mulhern, T. D. (2001). ATM, a central controller of cellular responses to DNA damage. Cell Death and Differentiation 8, 1052-1065.

39. Durocher, D. and Jackson, S. P. (2002). The FHA domain. Febs Letters 513, 58-66.

40. D'Amours, D. and Jackson, S. P. (2002). The Mre11 complex: at the crossroads of DNA repair and checkpoint signalling. Nature Reviews Molecular Cell Biology 3, 317-327.

41. Jazayeri, A. and Jackson, S. P. (2002). Screening the yeast genome for new DNA- repair genes. Genome Biology 3, reviews1009.1-1009.5.

42. Jackson, S. P. (2002). Sensing and repairing DNA double-strand breaks. Carcinogenesis 23, 687-696.

43. Rouse, J. and Jackson, S. P. (2002). Interfaces between the detection, signaling, and repair of DNA damage. Science 297, 547-51.

44. Bradbury, J. M. and Jackson, S. P. (2003). The complex matter of DNA double-strand break detection. Biochemical Society Transactions 31, 40-44.

45. Downs, J. A. and Jackson, S. P. (2003). Cancer: protective packaging for DNA. Nature 424, 732-734.

46. Gravel, S. and Jackson, S. P. (2003). Increased genome instability in aging yeast. Cell 115, 1-2.

47. Bradbury, J.M. and Jackson, S.P. Double-strand break recognition and its repair by nonhomologous end joining. In Handbook of Cell Signaling, Volume 3 p. 219-224 (Academic Press).

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50. Reaper, P. M., Di Fagagna, F. D. and Jackson, S. P. (2004). Activation of the DNA Damage Response by Telomere Attrition: A Passage to Cellular Senescence. Cell Cycle 3, 543-546.

51. Stucki, M. and Jackson, S. P. (2004). MDC1/NFBD1: a key regulator of the DNA damage response in higher eukaryotes. DNA Repair (Amst) 3, 953-957.

52. Downs, J. A. and Jackson, S. P. (2004). A means to a DNA end: the many roles of Ku. Nature Reviews Molecular Cell Biology 5, 367 -378.

53. d'Adda di Fagagna, F., Teo, S. H. and Jackson, S. P. (2004). Functional links between telomeres and proteins of the DNA-damage response. Genes and Development 18, 1781-1799.

54. Surralles, J., Jackson, S. P., Jasin, M., Kastan, M. B., West, S. C. and Joenje, H. (2004). Molecular cross-talk among chromosome fragility syndromes. Genes and Development 18, 1359-1370.

55. Stucki, M. and Jackson, S. P. (2004). Tudor domains track down DNA breaks. Nat Cell Biol 6, 1150-1152.

56. von Zglinicki, T., Saretzki, G., Ladhoff, J., Fagagna, F. and Jackson, S. P. (2005). Human cell senescence as a DNA damage response. Mech Ageing Dev 126, 111- 117.

57. -damage- response machinery to broken chromosomes. DNA Repair 5, 534-543.

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59. Jackson S.P. (2009). The DNA-damage response: new molecular insights and new approaches to cancer therapy. Biochem. Soc. Trans. 37, 483-94.

60. Giunta, S., and Jackson, S.P. (2011). Give me a break, but not in mitosis: The mitotic DNA damage response marks DNA double strand breaks with early signaling events. Cell Cycle, 10, 1215-1221.

61. Polo, S.E., and Jackson, S. P. (2011). Dynamics of DNA damage response proteins at DNA breaks: a focus on protein modifications. Genes and Development. 25, 409- 433.

62. Miller, K.M. and Jackson, S.P. (2012). Histone marks: repairing DNA breaks within the context of chromatin. Biochemical Society Transactions, 40(2), 370-376.

63. Forment, J.V., Kaidi, A. and Jackson, S.P. (2012). Chromothripsis and cancer: causes and consequences of chromosome shattering. Nature Reviews Cancer 12, 663-670.

64. Jackson, S.P*. and Durocher, D*. (2013). Regulation of DNA responses by Ubiquitin and SUMO. Molecular Cell 49, 795-807. *Co-corresponding authors.

65. Chahwan R., Gravel S., Matsusaka T., Jackson S.P. (2013). RNF8 links nucleosomal and cytoskeletal ubiquitylation of higher order protein structures. Cell Cycle 12, 1-5.

66. Schmidt, C. and Jackson, S.P. (2013). On your MARK, Get SET(D2), Go! H3K36me3 Primes DNA Mismatch Repair. Cell 153, 513-515.

67. Jacq, X., Kemp, M., Martin, N.M.B. and Jackson, S.P. (2013). Deubiquitylating enzymes and DNA damage response pathways. Cell Biochem Biophys 67, 25-43.

68. Jacq, X., Kemp, M., Martin, N.M. and Jackson, S.P. (2013). Deubiquitylating enzymes and DNA damage response pathways. Cell Biochem Biophys 67, 25-43.

69. Belotserkovskaya, R. and Jackson, S.P. (2014) Keeping 53BP1 out of focus in mitosis. Cell Research 37

70. Brown, J. S. and Jackson, S. P. (2015). Ubiquitylation, neddylation and the DNA damage response. Open Biology 5(4), 150018.

71. Beli P and Jackson SP. (2015) Ubiquitin regulates dissociation of DNA repair factors from chromatin. Oncotarget 6, 14727-14728.

APPENDIX: From Ku to KuDOS and the clinical concept of synthetic lethality

Two major factors led me to establish KuDOS Pharmaceuticals Ltd. and the drug discovery programmes within it. First, during the mid 1990’s, my group developed biochemical screening methods for DNA-PK and other DDR enzymes, leading us to identify initial small-molecule compounds that could inhibit these enzymes and enhance cancer cell killing by DNA damaging agents. Second, also during the mid 1990’s, our work in yeast led us to realize that defects in some DDR pathways only cause marked DNA-damage hypersensitivity in the absence of another DDR pathway. As explained below, such “synthetic lethal” relationships between DDR pathways can be exploited to striking clinical effect.

As outlined in the original KuDOS Business Plan that I wrote in 1997, because they lack particular DDR pathways, one might expect cancer cells to be more susceptible than normal cells to inhibition of certain other DDR pathways:

Although there are many examples of DDR impairment in cancer cells, a prime example is the loss of BRCA1 or BRCA2 function in hereditary breast and ovarian cancers. Here, the patient inherits one inactive copy of BRCA1/2, meaning that she (or he) has a high risk of developing cancer, particularly breast and ovarian malignancies. Notably, while normal cells in such a cancer patient possess a wild-type copy of the BRCA1/2 gene – rendering them BRCA1/2 proficient – the tumour cells have invariably lost the function of the other BRCA1/2 allele, making them dysfunctional for BRCA1/2. Crucially, work between 1997-2001 revealed that BRCA1/2 defects markedly impair DSB repair by HR. In light of these findings, in mid 2002 I (as KuDOS CSO) approached Alan Ashworth (Breakthrough Breast Cancer Centre, London, UK) with the proposal to assess the impact of DNA-PK, ATM and PARP-1 inhibitors in cells proficient or deficient in BRCA1 or BRCA2. Our ensuing collaborative work quickly revealed that BRCA-/- cells are strikingly hypersensitive to PARP inhibition, being killed at concentrations that are much, much lower than needed to kill BRCA+/+ or BRCA+/- cells. Parallel work by Thomas Helleday (University 38

of Sheffield, UK and University of Stockholm, Sweden) and Nicola Curtin (University of Newcastle-Upon-Tyne, UK) reached similar conclusions. These and subsequent findings thus established PARP inhibitors as the first of a new category of anti-cancer agents that operates on the principle of synthetic-lethality: targeting a pathway that is not essential to normal cells because they possess a compensatory pathway, but which is essential to cancer cells because they lack the compensatory pathway.

On the basis of the above findings and subsequent data generated with mouse cancer models, the orally-active KuDOS PARP inhibitor LynparzaTM/olaparib (formerly KU- 0059436/AZD2281), was selected by AstraZeneca for a clinical development. While trial- design and formulation issues slowed olaparib’s progress, recently received FDA and EMA approval for use in ovarian cancers with BRCA1/2 mutations. LynparzaTM is thus the first-in-class drug targeting PARP, and is the first drug to exploit the synthetic-lethality concept. It is currently being evaluated in seven Phase III trials.

LAUREATES OF THE GAGNA A. & CH. VAN HECK PRIZE

2003 : ► Michael L. J. APUZZO, University of Southern California, Los Angeles, USA

► Anne DEJEAN-ASSEMAT, Institut Pasteur, Paris, France and Laurent DEGOS, Hôpital St-Louis, Paris, France

2006 : ► Alexander J. VARSHAVSKY, California Institute of Technology, Pasadena, USA

2009 : ► Agnès RÖTIG, Hôpital Necker-Enfants Malades, Paris, France and Arnold MUNNICH, Hôpital Necker-Enfants Malades, Paris, France

2012 : ► Yehezkel BEN-ARI, InMed - INSERM, Marseille, France

2015 : ► Stephen P. JACKSON, The Gurdon Institute, University of Cambridge, UK

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