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Papers highlighted in the milestones section were chosen with the help of a panel of experts listed below:
Bruno Amati Jiri Bartek Robert Benezra Julian Blow
Bruce Bowerman Anthony Carr Ron DePinho John Diffley
Willian Dunphy Nick Dyson Bruce Edgar Stephen Elledge
Mark Ewen Iain Hagan M. Andrew Hoyt Tim Hunt
Lee Johnston Marc Kirschner Jurgen Knoblich Wilhelm Krek
Hartmut Land Nick La Thangue Christoph Lengauer Joachim Li
James Maller Yoshio Masui Clare McGowan Jonathan Millar
Timothy Mitchison David Morgan Kim Nasmyth Joseph Nevins
Eisuke Nishida Pat O'Farrell Moshe Oren Helen Piwnica-Worms
Gordon Peters Jan-Michael Peters Jonathon Pines Martin Raff
Steve Reed Jim Roberts Paul Russell Charles Sherr
Peter Sorger Bruce Stillman Mike Tyers Frank Uhlmann
Bert Vogelstein Karen Vousden Jean Wang Robert Weinberg
Mark Winey Curt Wittenberg Anthony Wynshaw-Boris Mitsuhiro Yanagida
 
Bruno Amati
Bruno Amati is a senior group leader at the DNAX Research Institute, Palo Alto, California. After graduating in Biology at the University of Geneva, Bruno pursued his Ph.D. studies with Susan Gasser at the Swiss Institute for Cancer Research (ISREC) on higher-order chromatin folding in yeast. In 1990, he joined the group of Hartmut Land at the Imperial Cancer Research Fund in London, where he initiated his research on the c-MYC oncoprotein. He returned to ISREC as a junior group leader in 1994 and joined DNAX in 1999. With MYC as the primary focus, research in Bruno's group has concentrated on molecular mechanisms of cell-cycle control, transcription and chromatin modifications.
 
Jiri Bartek
Jiri Bartek is the head of the Department of Cell Cycle and Cancer at the Institute of Tumour Biology of the Danish Cancer Society in Copenhagen, Denmark. He received his M.D. degree from Palacky University in Olomouc, and his Ph.D. degree in Cell Biology from the Institute of Molecular Genetics in Prague, Czech Republic. Prior to his present appointment, Jiri Bartek was a research group leader and department head in research institutes in Brno and Prague, Czech Republic, and spent several years as a visiting scientist at the Imperial Cancer Research Fund in London and the German Cancer Research Center in Heidelberg. His main research interests include the molecular mechanisms of mammalian cell-cycle control and responses to DNA damage, and the cancer-predisposing aberrations of these regulatory pathways.
 
Robert Benezra
Robert Benezra is a Member at the Memorial Sloan–Kettering Cancer Center in the Department of Cell Biology, and a Professor of Biology at Cornell Graduate School of Medical Sciences in New York City. Before he joined Sloan–Kettering, Robert Benezra worked at the Fred Hutchinson Cancer Center in Seattle, where he identified the Id proteins and showed that they are key regulators of tumour growth, angiogenesis and metastasis. Benezra and colleagues also identified the first human mitotic checkpoint gene, MAD2, and demonstrated its requirement for the execution of the checkpoint, its ability to act as a negative regulator of the anaphase-promoting complex and its role in maintaining genomic stability in embryonic and somatic cells. His programme continues to focus on the molecular basis of aneuploidy, tumour initiation and metastasis.
 
Julian Blow
Julian Blow is head of the Cancer Research Campaign Chromosome Replication Research Group in Dundee. He did his Ph.D. in Ron Laskey's laboratory in Cambridge in 1987, and then did postdoctoral research with Paul Nurse in Oxford. In 1991, he joined the Imperial Cancer Research Fund, before moving to the University of Dundee in 1997. His research interests are focused on cell-cycle control and the regulation of chromosome replication. He pioneered the use of Xenopus egg extracts for the study of chromosome replication and has developed the concept of replication licensing and its central role in co-ordinating DNA replication with cell-cycle progression.
 
Bruce Bowerman
Bruce Bowerman did his Ph.D. with Harold Varmus at the University of California in San Francisco. During this time, he developed an in vitro reaction for the integration of retroviral DNA into chromosomal DNA. He worked on Caenorhabditis elegans embryogenesis as a postdoctoral fellow with Jim Priess at the Fred Hutchinson Cancer Research Center in Seattle, from 1989–1992, before starting his own laboratory in the Institute of Molecular Biology at the University of Oregon, in Eugene, where he has been ever since. His laboratory aims to identify different pathways that regulate cell polarity and cell division in the early C. elegans embryo. Their goal is to understand how different pathways interact to execute asymmetric cell divisions that differentially segregate different developmental potential to daughter cells.
 
Anthony Carr
Anthony Carr studied for his B.Sc. at Norwich University, graduating in Biological Sciences in 1981. He subsequently studied under Paul Nurse for his D.Phil. at the University of Sussex. The subject of his thesis was "The analysis and function of the cdc2 gene region in fission yeast". Over the following years, he developed his interest in the checkpoint pathways that respond to DNA damage and replication delays, while working on DNA repair at the Medical Research Council (MRC) Cell Mutation Unit, Sussex University. His laboratory currently studies the genetics and biochemistry of these pathways in fission yeast, and they are interested in the mechanisms by which checkpoints respond to DNA damage and how these pathways interact with aspects of DNA repair and replication. His laboratory is now part of the Centre for Genome Damage and Stability, which is based at Sussex University and is funded by the MRC.
 
Ron DePinho
Ron DePinho is the American Cancer Society Research Professor and Professor of Medicine and Genetics at Harvard Medical School, Boston, USA. Before he joined the Dana–Farber Cancer Institute at Harvard, Ron DePinho was a professor at the Albert Einstein College of Medicine. His research has focused on key genetic events in the life-history of cancer cells, including those that impact on cell-cycle control (the p16–RB pathway), cellular survival (ARF–p53 pathway), cellular crisis (telomeres and telomerase) and host–tumour interactions. He pioneered the concept of tumour maintenance and the role of cellular crisis in cancer. His programme makes extensive use of engineered mouse models of human cancer to understand complex interactions between cancer-relevance pathways on the organismal level.
 
John Diffley
John Diffley was born and raised in the New York City area. He obtained his B.A., M.S. and Ph.D. from New York University. His Ph.D. dissertation was on the structure and mechanism of DNA polymerase alpha from Drosophila melanogaster. John then worked as a postdoctoral fellow in Bruce Stillman's laboratory at Cold Spring Harbor from 1984–1990, where he began working on yeast DNA replication. This work has continued in his laboratory at the Imperial Cancer Research Fund.
 
William Dunphy
William Dunphy is a professor in the Division of Biology at the California Institute of Technology and an Associate Investigator in the Howard Hughes Medical Institute. He received an A.B. degree in Biochemical Sciences from Harvard College and a Ph.D. degree in Biochemistry from Stanford University. He was a Helen Hay Whitney postdoctoral fellow at the University of California, San Diego. His honours include a scholar award from the Lucille P. Markey Charitable Trust and a presidential young investigator award from the National Science Foundation.
 
Nick Dyson
Nick Dyson is a member of the Massachusetts General Hospital Cancer Center and Associate Professor at Harvard Medical School, Boston. Nick received his Ph.D. from the University of Manchester in 1988 and he reached Boston by way of Amersham International and Cold Spring Harbor Laboratory. His laboratory studies cell-cycle control, and uses Drosophila as a model system to examine the functions of Rb- and E2F-family members.
 
Bruce Edgar
Bruce Edgar is a member of the basic sciences division as the Fred Hutchinson Cancer Research Center (FHCRC) in Seattle, Washington. Bruce graduated from Swarthmore College in Pennsylvania and conducted his postgraduate degree in the laboratory of Gerold Schubiger at the University of Washington, Seattle, while studying early embryonic development in Drosophila melanogaster. During his postdoctoral work in the laboratory of Patrick O'Farrell at the University of San Francisco, Bruce turned his attention to understanding cell-cycle regulation in Drosophila. After a brief period working in the laboratories of Paul Nurse and Phil Ingham at the Imperial Cancer Research Fund in London and looking at cell-cycle control in fission yeast, Bruce returned to studying the cell cycle in Drosophila. Since 1993, Bruce has been based at the FHCRC where he has continued his work on understanding the genetics behind the cell-cycle programme in Drosophila.
 
Stephen Elledge
Steve Elledge is a professor with the Howard Hughes Medical Institute and is the Robert A. Welch Professor of Biochemistry in the Department of Biochemistry and Molecular Biology at the Baylor College of Medicine, Texas. He received his Ph.D. degree in Biology from the Massachusetts Institute of Technology, Boston. He was a Helen Hay Whitney postdoctoral fellow at Stanford University, California. Steve's current research interests are in understanding how cells sense and respond to DNA damage and the development of new genetic technologies to facilitate mammalian genetics. Steve Elledge is a Pew Scholar in the biomedical sciences. He recently received the G.H.A. Clowes Memorial Award and the Michael E. DeBakey Award for research excellence.
 
Mark Ewen
Mark Ewen holds appointments in the Department of Adult Oncology at the Dana–Farber Cancer Institute and the Department of Medicine at Harvard Medical School. Mark and his collaborators demonstrated that the retinoblastoma susceptibility gene product (RB) was a critical target of D-type cyclin action, and thereby contributed to delineation of the RB pathway. His work also includes the cloning of the gene for the RB-related protein p107, and the discovery that a significant proportion of RAS-mediated mitogenic signalling is exerted through cyclin D1, KIP1 and thereby RB. These efforts have contributed to our understanding of the function of critical cell-cycle control elements and how their deregulation can contribute to human cancer. Current studies are devoted to understanding the oncogenic actions of cyclin D1 and the genetic interactions between RB and RAS.
 
Iain Hagan
Iain Hagan has a chair in Cell Biology at the University of Manchester and his laboratory is based at the Paterson Institute for Cancer Research in Manchester. After a Ph.D. under the supervision of Jerry Hyams (University College London) and Paul Nurse (Imperial Cancer Research Fund), he spent a brief spell doing postdoctoral research in Paul Nurse's laboratory in Oxford, before undertaking a second postdoctoral position in Mitsuhiro Yanagida's laboratory in Kyoto, Japan. After four years in Kyoto, he returned to the UK with a Cancer Research Campaign return fellowship to set up a laboratory in the School of Biological Sciences in the University of Manchester. Throughout his career, he has used the fission yeast Schizosaccharomyces pombe as a model system to study the mechanics and control of cell division.
 
M. Andrew Hoyt
Andrew Hoyt received his Ph.D. from University of California in Berkeley, where he performed molecular genetic studies of phage development under the tutelage of Hatch Echols. As a postdoctoral fellow with David Botstein at the Massachusetts Institute of Technology, he learned to apply genetic techniques to problems in eukaryotic cell biology. Andrew Hoyt moved to The Johns Hopkins University in 1988, where he is currently a Professor of Biology. Using the yeast Saccharomyces cerevisiae for a model organism, his laboratory studies the mechanisms and regulation of mitotic cell division. Determining the mitotic roles performed by microtubule-based motor proteins has been a major focus. Also, inspired by the cell-cycle checkpoint concept of Ted Weinert and Leland Hartwell, the Hoyt laboratory identified and characterized key mitotic-spindle-checkpoint regulatory proteins.
 
Tim Hunt
Tim Hunt started his scientific career in 1964 in the Department of Biochemistry at Cambridge, under the supervision of Asher Korner, where he worked on the control of haemoglobin synthesis. He then moved to New York City to do postdoctoral research with Irving London, where he pursued this interest. Tim returned to Cambridge, but took the opportunity of teaching summer courses at the Marine Biological Laboratory, Woods Hole, so he could look at changes in protein synthesis in sea urchin and clam eggs after fertilization. This work led to the discovery of cyclins — proteins that oscillate throughout the cell cycle — which bind and activate cyclin-dependent kinases. Tim Hunt, Leland Hartwell and Paul Nurse were awarded the 2001 Nobel Prize for Physiology and Medicine for the discovery of cyclins and cyclin-dependent kinases. Tim moved to the Imperial Cancer Research Fund in 1991, where he has continued this work.
 
Lee Johnston
Lee Johnston is Head of the Division of Yeast Genetics at the National Institute for Medical Research in Mill Hill, London. He has investigated various aspects of yeast cell-cycle control, initially focusing on DNA replication and control of S phase. The latter work included the identification of a transcriptional system for co-expression of genes required for S phase, a system with close analogies to E2F–DP1 in mammalian cells. Extending this work revealed an interaction between cell-cycle-dependent transcription and the Pkc1-dependent MAP kinase pathway to coordinate events in late G1 phase. Subsequently, his laboratory focused on control of mitotic exit and identified the mitotic exit network and a role for the CDK inhibitor Sic1 in mitotic kinase inactivation.
 
Marc Kirschner
Marc Kirschner began to work seriously on the cell cycle after experiments with Koki Hara in 1978 showed that an enucleated frog egg underwent periodic contractions of its cortex that timed precisely with the cell cycle of cleaving eggs. Later, in collaboration with John Gerhart, he showed that this enucleated cycle coincided with periodic oscillations in maturation-promoting factor (MPF), and that MPF activity drove events such as spindle assembly, nuclear envelope breakdown and DNA replication. With Martha Cyert and Andrew Murray, Marc developed an in vitro system that carried out the entire embryonic cell cycle. They used this to show that cyclin was the only protein whose synthesis was required to drive the cell cycle and to describe the role of phosphorylation in regulating cdc2. They also showed that cyclin degradation was required for cell-cycle exit, and that this occurred through its ubiquitylation. Marc and colleagues characterized and purified the anaphase-promoting complex (APC), which targets cyclin for ubiquitylation, and than helped to describe how APC leads to chromatid separation through the degradation of securin, and how the checkpoint blocks APC through MAD2. Most recently they have described the regulation of separase by phosphorylation and degradation.

Photo Stu Rosner Copyright 2000, PO Box 397, Cambridge MA 02140, Phone: 617-576-1897, Fax: 617-576-6191, All rights reserved
 
Jurgen Knoblich
Juergen Knoblich did his Ph.D. on cell-cycle control in Drosophila with Christian Lehner at the Max Planck Institute in T�bingen. For his postdoctoral studies, he joined Yuh Nung Jan's laboratory at the University of California, San Francisco, to work on asymmetric cell division in Drosophila. He is now a group leader at the Research Institute of Molecular Pathology in Vienna, where his group is studying how asymmetric cell divisions are orientated, and how cell-fate determinants are segregated into one of the two daughter cells.
 
Wilhelm Krek
Wilhelm Krek is Senior Group Leader and Coordinator of the Growth Control Program at the Friedrich Miescher Institute for Biomedical Research in Basel. After studying chemistry at the University of Graz, Wilhelm did his Ph.D. research with Erich Nigg on cell-cycle-control mechanisms at the Swiss Cancer Institute in Lausanne. He then moved to the Dana–Farber Cancer Institute in Boston, as a postdoctoral fellow, to work with David Livingston on E2F transcription factors and their role in cancer. In 1995, Wilhelm returned to Switzerland as a Junior Group Leader at the Friedrich Miescher Institute for Biomedical Research. Wilhelm Krek's laboratory is interested in the molecular mechansims of tumour-cell evolution.
 
Hartmut Land
Hartmut Land is the Robert and Dorothy Markin Professor and Chair of the Department for Biomedical Genetics and the Director of Basic Sciences in the James P. Wilmot Cancer Center at the University of Rochester, New York. Having received his Ph.D. in Biology from the University of Heidelberg, he undertook postdoctoral research at the Massachusetts Institute of Technology and the Whitehead Institute. In 1985 he moved to the Imperial Cancer Research Fund, where he headed the Growth Control and Development Laboratory. He has been at the University of Rochester since 1999. His research interests are the molecular and mechanistic basis of multistage carcinogenesis.
 
Nick La Thangue
Appointed to the Cathcart Chair of Biochemistry, University of Glasgow in 1995, Nick La Thangue's research interests have focused on the role of the retinoblastoma tumour suppressor protein (RB) and E2F in regulation of the G1 to S phase transition. During his career, and latterly at the UK Medical Research Council National Institute for Medical Research, his work uncovered the regulation of E2F by RB, together with the role of viral oncoproteins, including the adenovirus E1A protein, in targeting the RB–E2F complex. More recently, Nick's research has investigated the molecular mechanisms through which E2F promotes cell-cycle progression.
 
Christoph Lengauer
Christoph Lengauer is Assistant Professor of Oncology at the Johns Hopkins University School of Medicine and Director of the Cell Imaging Core Facility of the Johns Hopkins Oncology Center. He holds an M.B.A. degree in Medical Services Management and received his Ph.D. degree in Biology from the University of Heidelberg, Germany, where he developed multicolour fluorescence in situ hybridization protocols for the staining of chromosomal material. He was a Boehringer Ingelheim Fonds postdoctoral fellow at the Institute for Molecular Pathology in Vienna, Austria and an Erwin–Schroedinger postdoctoral fellow in the laboratory of Bert Vogelstein in Baltimore, Maryland. Over the past few years, Christoph Lengauer became interested in the problem of genetic instability and the mystery of aneuploidy in cancer. He has focused his research on cell-cycle regulation, checkpoint control and the mechanisms that maintain genomic integrity.
 
Joachim Li
Joachim Li graduated from the University of Chicago with a B.S. and M.S. in Chemistry in 1980. He received his Ph.D. and M.D. in 1988 from The Johns Hopkins University School of Medicine. During this period he developed and characterized an in vitro system for the replication of SV40 DNA in the laboratory of Thomas J. Kelly. Joachim was a postdoctoral fellow in the laboratory of Ira Herskowitz at the University of California, San Francisco (UCSF) from 1989–1993. In the Herskowitz laboratory, Joachim developed the one-hybrid assay to identify and clone proteins that bind yeast origins. In 1993, Joachim joined the faculty at UCSF, where he was named a Markey Scholar, a Searle Scholar and a Rita Allen Scholar. He is currently an Associate Professor in the Department of Microbiology and Immunology at UCSF. His laboratory studies the mechanism and cell-cycle regulation of replication initiation in budding yeast and is investigating the consequences of disrupting this regulation to genomic stability.
 
James Maller
James Maller is an Investigator of the Howard Hughes Medical Institute and a Professor of Pharmacology at the University of Colorado School of Medicine, Denver, USA. He received his Ph.D. in Molecular Biology from the University of California, Berkeley, where he worked with John Gerhart and carried out postdoctoral work with Edwin Krebs at the University of Washington. He pioneered research on the cell cycle that first purified mitosis-promoting factor (MPF) and then identified its subunits as products of the Cdc2 and cyclin B genes. His laboratory continues to unravel multiple protein kinase cascades that control entry into and exit from mitosis.
 
Yoshio Masui
Yoshio Masui earned his Ph.D. at Kyoto University, Japan, in 1961, and became Assistant Professor at Konan University, Japan, in 1965. In 1966, he went to Yale University, USA, on his sabbatical leave to study oocyte maturation. He moved to the University of Toronto, Canada, in 1969 as Associate Professor, and was appointed to Professor in 1978. He retired in 1997 and is now Professor Emeritus at the University of Toronto.
 
Clare McGowan
Clare McGowan did her Ph.D., studying protein phosphatases, with Phillip Cohen in Dundee. For her postdoctoral training, she went to Paul Russell's laboratory, where she began to apply genetic advances made in fission yeast to the identification and characterization of human cell-cycle regulators. Despite her plan to stay there for just two years, Clare was seduced by the sea, sun and science, and she took a faculty position at The Scripps Research Institute, La Jolla, California. The guiding principle of the laboratory remains the translation of genetic leads from model organisms to analysis of relevant gene products in human cells.
 
Jonathan Millar
Jonathan Millar is a Senior Staff Scientist at the National Institute for Medical Research (NIMR) in London. While a postdoctoral scientist in Paul Russell's laboratory at The Scripps Research Institute, La Jolla, California, he showed that the CDC25 gene product encodes a specialized phosphatase that triggers mitotic initiation in all eukaryotes. He and his colleagues have continued to study the signalling pathways that govern cell-cycle progression in fission yeast and have recently uncovered a new mitotic checkpoint that prevents the separation of sister chromatids when spindles are misoriented. Their current studies are devoted to understanding the controls that ensure equal segregation and accurate duplication of chromosomes in each cell cycle.
 
Timothy Mitchison
Timothy Mitchison is a Professor of Cell Biology and co-director of the Institute of Chemistry and Cell Biology at Harvard Medical School, Boston, USA. Before he joined Harvard, Tim was a Professor of Pharmacology at the University of California, San Francisco. His current research focuses on microtubule and actin dynamics, and how these contribute to cell division and cell migration. He is also interested in synthesis and discovery of small-molecule tools for basic cell biology, and in development of microscope-imaging technology.
 
David Morgan
David Morgan studies the molecular mechanisms that govern progression through the eukaryotic cell cycle, with an emphasis on the regulation and targets of the cyclin-dependent kinases. He is a Professor of Physiology and Biochemistry at the University of California in San Francisco (UCSF), USA. After an undergraduate degree in Animal Physiology at the University of Calgary, David moved to UCSF to study insulin and growth-factor-receptor signalling with Richard Roth for his Ph.D., and with William Rutter as a postdoctoral fellow. His interest in cell-cycle control developed during a second postdoctoral position, with Harold Varmus at UCSF.
 
Kim Nasmyth
Born in London in 1952, Kim Nasmyth was educated in Britain and obtained his Ph.D. degree in Zoology from the University of Edinburgh. For his doctoral thesis, completed in 1977, he chose the topic of DNA replication in yeast. In 1988, after several years of research in the USA and in Cambridge, Kim joined the newly established Research Institute of Molecular Pathology in Vienna. At present, he is its Managing Director. Kim continues to be very active in research, using yeast to study mechanisms concerned with chromosome segregation. Recent breakthroughs include the identification of proteins that hold sister chromatids together and a protease that splits them apart. Kim Nasmyth has been awarded numerous scientific honours, including the Max Perutz Prize, the Unilever Science Prize, the Louis Jeantet Prize for Medicine and the Wittgenstein Prize. Kim Nasmyth is a Fellow of the Royal Society.
 
Joseph Nevins
Joseph Nevins is James B. Duke Professor and Chair of the Department of Genetics at Duke University Medical Center. He is also an Investigator with the Howard Hughes Medical Institute. He and his colleagues discovered the E2F cellular transcription factor and then identified E2F as a functional target for the RB tumour-suppressor protein. Further work by Nevins and others has defined the role of E2F in the control of genes that are critical for S phase, so providing a mechanistic basis for the growth-regulatory role of RB.
 
Eisuke Nishida
Eisuke Nishida is a Professor at Kyoto University. Before he joined Kyoto University in 1993, Eisuke Nishida was at the University of Tokyo where he and his colleagues found a serine/threonine kinase activity towards MAP2 in vitro that was commonly activated by growth factors and tumour promoters (now called MAP kinase). They also identified two mammalian MAP kinase polypeptides, corresponding to p44MAPK (ERK1) and p42MAPK (ERK2). They then identified a direct activator of MAP kinase (MAP kinase kinase, MEK) and played the leading part in establishing the concept of the MAP kinase cascade. Their research has focused on regulatory mechanisms and functions of the MAP kinase cascade and other signal-transduction pathways in the cell cycle, cell proliferation, differentiation and developmental processes.
 
Pat O'Farrell
As a student, Pat O'Farrell developed the technique of two-dimensional gel electrophoresis, and a global approach to problems that is enjoying new life under the name of 'proteomics'. His laboratory, in the Department of Biochemistry and Biophysics at the University of California in San Francisco (UCSF), has studied embryonic pattern formation in Drosophila. In one of the first applications of positional cloning, they identified the engrailed segmentation gene, and demonstrated its striped expression. They showed that homeodomain regulators bind specific DNA sequences and modulate transcription, and that Wingless transmits patterning signals from one cell to another. Their studies of cell-cycle control in embryos showed that developmentally programmed transcription of a mitotic activator controls where and when cells divide. Present work fuses genetic and cell-biological approaches in analyses of mitotic events, and new efforts explore regulation of innate immunity in Drosophila.
 
Moshe Oren
Moshe Oren received his Ph.D. from the Weizmann Institute in 1978. In 1981, he returned to the Weizmann Institute as a faculty member. In 1983, in collaboration with Arnold Levine in Princeton, with whom he carried out his postdoctoral research, Moshe reported the cloning of the gene that encodes the p53 tumour-suppressor protein. He has continued to be active in the p53 field, and has made many contributions to the understanding of the mode of action and regulation of p53. Moshe is currently Dean of the Faculty of Biology at the Weizmann Institute.
 
Helen Piwnica-Worms
Helen Piwnica-Worms is Professor of Cell Biology and Physiology and Professor of Medicine at Washington University School of Medicine, St. Louis. She is also an Investigator of the Howard Hughes Medical Institute. Helen and her colleagues study how entry into mitosis is regulated during a normal cell cycle; how entry into mitosis is prevented when unreplicated and/or damaged DNA is detected; and how cell-cycle and checkpoint pathways are disrupted in human cancers. In addition to showing how the mitotic inducer CDC2 is regulated, Helen and her group discovered pathways linking G2 checkpoint regulators to the mitotic inducer CDC25C. These pathways involve 14-3-3 proteins and the checkpoint kinases ATM, ATR, CHK1 and CHK2. Helen is interested in anticancer agents that target these regulators of G2 checkpoint control because they might prove useful when combined with DNA-damaging agents in treating p53-deficient human cancers.
 
Gordon Peters
Gordon Peters received a Ph.D. degree from the University of Edinburgh and, after postdoctoral training at the University of Wisconsin, joined the Imperial Cancer Research Fund (ICRF) in London in 1977. His early work on the identification and characterization of oncogenes activated by mouse mammary tumour virus and, subsequently, of gene amplification in human breast cancer, led to an interest in the dysregulation of cyclin-dependent kinases and their inhibitors in human tumours. He is currently a Principal Scientist at the ICRF and heads a laboratory working on the role of cell-cycle regulatory proteins and tumour suppressors in telomere-based and oncogene-induced senescence.
 
Jan-Michael Peters
Jan-Michael Peters' main interest is to obtain insight into the mechanisms and the regulation of mitosis in vertebrate cells. He is particularly interested in understanding how mitosis is controlled by proteolysis, which is mediated by the ubiquitin ligase APC (anaphase-promoting complex), and the protease separase. Jan obtained his Ph.D. in Biology from the University of Heidelberg in 1991, did postdoctoral research with Werner Franke in Heidelberg and Marc Kirschner in Boston, and has been a Group Leader at the Research Institute of Molecular Pathology in Vienna since 1996.
 
Jonathon Pines
Jonathon Pines has been attempting to understand the cell cycle, and in particular mitosis, since graduation. It all began as a Ph.D. student with Tim Hunt with whom he cloned sea-urchin cyclin B, and continued as a postdoctoral researcher with Tony Hunter, with whom he cloned human cyclins A and B1. He returned to the UK and set up a group at the Wellcome/Cancer Research Campaign Institute in Cambridge to continue working on mammalian mitosis. Recently, he has been seduced by the movie industry and is using GFP-fusion proteins to understand how proteins are targeted to specific subcellular structures during the cell cycle, and to analyse the role of proteolysis in coordinating mitosis.
 
Martin Raff
Martin Raff was born and educated in Montreal. He received his B.Sc. and M.D. degrees at McGill Universtiy and did a residency in medicine at the Royal Victoria Hospital in Montreal and in neurology at the Massachusetts General Hospital in Boston. He did postdoctoral training in immunology at the National Institute for Medical Research in London, after which he moved to University College London, where he has been a Professor of Biology since 1979. He is currently at the Medical Research Council Laboratory for Molecular Cell Biology and Cell Biology Unit and in the Biology Department at University College London. His research has been in immunology, cell biology, and developmental neurobiology. He is a Fellow of the Royal Society and the Academia Europaea, a foreign member of the American Academy of Arts and Sciences, was president of the British Society of Cell Biology from 1991 to 1995, and chairman of the UK Life Sciences Committee from 1998 to 2001.
 
Steve Reed
Steve Reed's primary research interest has always been to understand how cell proliferation is regulated. As a Ph.D. student at Stanford in the 1970s, he was involved in some of the earliest investigations of how DNA tumour viruses promote host cell-cycle transitions. Steve then moved to Leland Hartwell's laboratory to carry out postdoctoral research on yeast cell-cycle genes. He has continued this work ever since, first on the faculty of University of California in Santa Barbara, and currently as a professor at the Scripps Research Institute in La Jolla, California. About 10 years ago, his research group began to extrapolate the findings made in yeast to mammalian cells, which has turned out to be a highly productive research strategy. Currently, his laboratory maintains an active research programme focused on the cell cycle, and evenly split between yeast and mammalian systems.
 
Jim Roberts
Jim Roberts is a member of the Division of Basic Sciences at the Fred Hutchinson Cancer Research Center and an Investigator of the Howard Hughes Medical Institute. His work in the cell-cycle field began with experiments showing that CDK activation is a crucial step in starting S phase in mammalian cells. This led him to explore how CDK activity is controlled during the G1 to S phase transition. These experiments resulted in the discovery of cyclin E and then, in collaboration with Joan Massague, the discovery of the CDK inhibitor KIP1. His current work is focused on understanding the function and regulation of cyclin E and p27 in normal cells, and on understanding how these pathways are disrupted in human tumours.
 
Paul Russell
Paul Russell's laboratory uses the fission yeast Schizosaccharomyces pombe to study cell-cycle control, DNA-integrity checkpoints and cytotoxic stress responses. In recent years, his laboratory showed that DNA damage and replication checkpoints delay mitosis by regulating Cdc25 and Mik1, proteins that control the inhibitory tyrosine phosphorylation of Cdc2. Paul's interest in mitotic control stems from his postdoctoral studies with Paul Nurse at the Imperial Cancer Research Fund Laboratories in London. He received his Ph.D. in Genetics from the University of Washington, Seattle. He is currently a Professor of Molecular Biology at The Scripps Research Institute, La Jolla, California.
 
Charles Sherr
Charles Sherr is an Investigator of the Howard Hughes Medical Institute and Herrick Foundation Chairman of the Department of Tumor Cell Biology at St. Jude Children's Research Hospital in Memphis, Tennessee. He received his M.D. and Ph.D. degrees from New York University, and after completing an internship, worked for 10 years at the National Cancer Institute, before assuming his present position in 1983. Chuck is a cancer biologist whose main interests have focused on the roles of oncogenes and tumour suppressors, particularly as they affect progression through the mammalian cell-division cycle. He has received the Milken and Dameshek Prizes, the Bristol Myers–Squibb Award, and the Pezcoller Prize for his work in cancer research, and is a member of the U.S. National Academy of Sciences.
 
Peter Sorger
Peter Sorger, Ph.D., is an Associate Professor in the Department of Biology at the Massachusetts Institute of Technology (MIT). He also holds a joint appointment in the MIT Division of Bioengineering and MIT Center for Cancer Research, and is an affiliate of the Institute of Chemistry and Cell Biology at Harvard Medical School. Peter received his Ph.D. with Hugh Pelham at the Medical Research Council in Cambridge, UK, and postdoctoral training with Harold Varmus and Andrew Murray at the University of California in San Francisco. Sorger's research focuses on the mechanisms of chromosome–microtubule attachment and on the regulation of the spindle-assembly checkpoint in yeast and animal cells.
 
Bruce Stillman
Bruce Stillman is Director and Chief Executive Officer of Cold Spring Harbor Laboratory (CSHL). A native of Australia, he moved to Cold Spring Harbor as a postdoctoral fellow in 1979. Bruce has been Director of the CSHL Cancer Center since 1992, and in 1994 he succeeded James Watson as Director of CSHL. Bruce's research focuses on the mechanism and regulation of inheritance of both DNA and chromatin structures in eukaryotic cells, particularly in yeast and human cells. He is a Fellow of The Royal Society and a foreign member of the US National Academy of Sciences.
 
Mike Tyers
Mike Tyers is a Senior Scientist at the Samuel Lunenfeld Research Institute of Mount Sinai Hospital, Toronto, and Professor in the Department of Medical Genetics and Microbiology at the University of Toronto, where he holds a Canada Research Chair in Biochemistry. His laboratory uses biochemical and genetic approaches in yeast and mammalian cells to understand the mechanics of cell division. Current research includes SCF-dependent degradation pathways that control the G1/S transition, genome stability and gene expression, with emphasis on mechanisms of phosphorylation-dependent substrate recognition.
 
Frank Uhlmann
Frank Uhlmann studied Biochemistry at the University of T�bingen and worked with Hans Probst on the control of cellular DNA replication. He then moved to New York to do his Ph.D. with Jerard Hurwitz at the Memorial Sloan–Kettering Cancer Center on the enzymology of human replication factor C. As a postdoctoral fellow, he joined the laboratory of Kim Nasmyth at the Research Institute of Molecular Pathology in Vienna, where he discovered the protease separase that cleaves cohesin to trigger chromosome separation in anaphase. Since August 2000, Frank has been head of the Chromosome Segregation Laboratory at the Imperial Cancer Research Fund in London.
 
Bert Vogelstein
Bert Vogelstein and his colleagues have shown that colorectal tumours result from the gradual accumulation of genetic alterations in specific oncogenes and tumour-suppressor genes. They have shown that naturally occurring mutations in genes such as TP53 and APC can provide critical clues to their biochemical and physiological functions. This work has had substantial implications for diagnosis and treatment and has formed a model for understanding human tumorigenesis in general. Their current studies are devoted to further understanding the genes and pathways through which colorectal neoplasia develops, as well as to the translation of this knowledge to patient management.
 
Karen Vousden
Karen Vousden received her Ph.D. in Genetics from the University of London. She carried out postdoctoral fellowships with Chris Marshall at the Institute of Cancer Research in London and Douglas Lowy at the National Cancer Institute (NCI), before becoming Head of the Human Papillomavirus Group at the Ludwig Institute for Cancer Research in London. She left to join the ABL-Basic Research Program as Head of the Molecular Carcinogenesis Section in 1995, and is now Chief of the Regulation of Cell Growth Laboratory (RCGL), Center for Cancer Research, NCI. Her main research interests are the regulation and function of the p53 tumour-suppressor protein.
 
Jean Wang
Jean Wang is the Herbert Stern Professor of Biology at the University of California, San Diego. Her research has been directed at understanding the control of mammalian cell proliferation with a focus on the functional interaction between oncogenes and tumour-suppressor genes. Beginning with the c-ABL tyrosine kinase, her work has revealed a complex network that links ABL with the retinoblastoma tumour-suppressor protein RB, the mismatch-repair proteins, the ATM kinase, and p73 in the regulation of cell-cycle checkpoints and apoptosis. Their current studies address how this regulatory network governs the choice of normal progenitor cells to proliferate, differentiate or die — choices that are invariably compromised in cancer cells.
 
Robert Weinberg
Robert Weinberg's laboratory has been interested in the contributions of oncogenes, tumour-suppressor genes and telomerase to the cell-transformation process. In 1982, his laboratory discovered a point mutation in RAS, which represented the first somatic mutation documented in a human tumour-cell genome. In 1986, workers in his laboratory isolated the retinoblastoma tumour-suppressor gene, RB. More recently, the Weinberg laboratory has isolated TERT, the gene that encodes the catalytic subunit of human telomerase, and this has allowed them to transform normal human cells into tumour cells. At present, the laboratory is focusing on understanding the interactions between cancer cells and normal cells that have been recruited into the tumour mass and that help transformed tumour cells to proliferate.
 
Mark Winey
Mark Winey's laboratory studies the cell-cycle-controlled duplication of centrosomes in yeast and in vertebrate cells. He did his postdoctoral training in yeast cell biology with Breck Byers at the University of Washington. While there, he identified regulators of yeast centrosome duplication that served as the basis of his laboratory's work in the Department of Molecular, Cellular and Developmental Biology at the University of Colorado, Boulder. Their work has also touched on the spindle-assembly checkpoint and on nuclear pore complexes. He has also enjoyed collaborating with the Boulder Laboratory for 3D Fine Structure to analyse yeast morphology using advanced electron-microscopy techniques.
 
Curt Wittenberg
Curt Wittenberg is interested in mechanisms regulating cell cycle initiation in the buddingyeast. His interest in cell cycle regulation began as a postdoctoral fellow in Steve Reed�s laboratory where he participated in the discovery and characterization of yeast G1 cyclins. His laboratory has focused upon the regulation of gene expression, function and proteolysis of G1 cyclins. He is a member of the Departments of Molecular Biology and Cell Biology at The Scripps Research Institute in La Jolla, California.
 
Anthony Wynshaw-Boris
Anthony Wynshaw-Boris received his M.D. and Ph.D. degrees from Case Western Reserve University, followed by a clinical genetics and postdoctoral fellowship at the Harvard Medical School. He was head of the Mouse Models Unit in the National Human Genome Research Institute at the National Institutes of Health for 5 years, before moving to the Departments of Paediatrics and Medicine at the University of California in San Diego School of Medicine. His laboratory studies mouse models of human neurogenetic diseases and cancer.
 
Mitsuhiro Yanagida
Mitsuhiro Yanagida was born in Tokyo, Japan. After studying biochemistry at the University of Tokyo, he moved to Geneva, Switzerland, and studied the molecular biology of bacteriophage T4. He continued to study head morphogenesis of T4 as a junior faculty of Kyoto University, and shifted to chromosome biology and cell-cycle research using fission yeast Schizosaccharomyces pombe after he was promoted to a full professor in 1977. He is currently a dean of Graduate School of Biostudies. The research interests of his group are focused on inheritance mechanisms of chromosomes and cell-cycle regulations of chromosome dynamics.